Records of the Western Australian Museum

ISSN 0312 3162

Records

of the

Western Australian
Museum

Volume 22 Part 1 2003

i

Records

of the

Western Australian Museum

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Printed and published by the Western Australian Museum
© Western Australian Museum, December 2003
ISSN 0312 3162

Cover: Volutoconus hargreavesi calcarelliformis (Wells and Morrison, 2003), a new subspecies of volute
from the deep waters off northwestern Australia

Illustration by Jill Ruse.

Records of the Western Australian Museum 22: 1-7 (2003).

8 JAN 2004

The 3-dimensional anatomy of the North-Western Marsupial Mole
( Notoryctes canrinus Thomas 1920) using computed tomography.
X-ray and magnetic resonance imaging

Natalie Warburton 1 , Christopher Wood 2 , Christopher Lloyd 2 , Swithin Song 2 and Philip Withers 1

1 Department of Zoology, University of Western Australia
Crawley, Western Australia, 6009. E-mail: nwarbo@cyllene.uwa.edu.au
2 Department of Radiology, Royal Perth Hospital
Wellington Street, Perth, Western Australia, 6000

Abstract - The 3-D skeletal images obtained from reconstruction of CT scans
and X-rays, and soft-tissue images produced by MRI, provide invaluable
information of the internal and gross anatomy of the north-western marsupial
mole ( Notoryctes caurinus).

The conical skull, which is quite thin-walled dorsally and anteriorly but
strong in the basicranial region, has little evidence of the orbit or zygomatic
arch, and the smoothly-curved posterior region has no sagittal or occipital
crests. The vertebral column is remarkably strengthened, and in lateral view
has an unusual flat-shape. The cervical vertebrae appear to be greatly
compressed; 4 or 5 are completely fused (which is unique among marsupials).
The thoracic vertebrae are fairly robust with large neural spines. The lumbar
vertebrae are distinct, becoming large posteriorly towards the pelvis. The
sacral vertebrae are greatly expanded in size and are fused with the pelvis.
Particularly in the middle of the tail, the caudal vertebrae are greatly
developed, with large transverse processes and chevron bones. The pectoral
girdle is very anterior, with the shoulder articulation level with the anterior
cervical vertebrae just behind the skull, and low on the side of the body. The
humerus is robust, and the radius and ulna are very short. The bones of the
pelvis are highly derived, and fused to sacral vertebrae. The epipubic bones
are small and not ossified. An ossified patella is present and it has an unusual
large triangular keel.

The most apparent soft-tissue structure by MRI is a large amount of
subcutaneous fat, particularly around the ventral surface of the pelvis but also
dorsal to the pelvis and anteriorly around the shoulders. The major muscle
groups are visible, but distinction between individual muscles is not possible
except for the very large muscles of the thigh, upper arm and base of the tail.
The muscles of the tail are strongly developed, more so ventrally than
dorsally.

INTRODUCTION

Computed tomography (CT scan) and magnetic
resonance imaging (MRI) are used extensively in
medical diagnostics and research and the
techniques have also been incorporated into the
veterinary discipline in clinical situations.
However, the zoological use of such technology
in more basic anatomical research has been rather
limited. The few anatomical investigations
incorporating CT and MRI technology include
descriptions of the internal anatomy of the feline
abdomen (Samii et al. 1999), and the brain and
coelomic cavity of the domestic pigeon
(Romagnano et al. 1996). The techniques have also
been used in paleaontological studies, for
example the measurements of brain size in
Allosaurus (Rogers 1999). This research uses CT

scan and MRI techniques to investigate the
anatomy of the marsupial mole.

The anatomy of the North-western marsupial
mole, Notoryctes caurinus (Marsupialia:
Notoryctidae), has not been studied in any detail
since its original description by Thomas (1920). In
fact the only published research for this remarkable
fossorial marsupial since then is the physiological
study of Withers et al. (2000) and a natural history
account by Thompson et al. (2000-2001). Very few
preserved specimens of N. caurinus are available, so
it is important to glean as much anatomical
information as non-destructively as possible from
the few specimens that are available. The purpose
of this paper is to provide an account of the
usefulness of CT scan, X-ray, and MRI to describe
N. caurinus , a small (mass - 30 - 35 g) and highly
specialized marsupial.

2

N. Warburton, C. Wood, C. Lloyd, S. Song, P.C. Withers

METHODS

The primary specimen of N. caurinus which we
examined was from the Western Australian
Museum (specimen number M 44175). Capture
locality was Cotton Creek (Newman-Canning Stock
Route road; 4/3/1999; 22°59' 122°23’). The specimen
was formalin fixed and preserved in 70% ethanol.
The specimen was male, with a total body length of
10.5 cm and mass of approximately 35 g.

CT and MRI scans were performed at Royal Perth
Hospital, Perth, Western Australia. The CT scans
were made with on a GE Cti scanner (General
Electric Medical Systems, Milwaukee, USA) using a
helical technique with a slice collimation of 1 mm,
and a pitch of 1:1. Images were reconstructed at an
interval of 0.3 mm using soft tissue and bone
algorithms. 3-D volume rendered images were also
obtained using the AW 3.0 workstation.

MRI scans were performed using a Marconi 1.0T
MR scanner (Marconi Medical Systems, Highland
Heights, Ohio, USA) using a 3-D RF FAST sequence
with 1.4 mm slices, flip angle of 35 degrees, TR of
24 ms, TE of 6 ms, field of view of 12 cm, 192x256
matrix, one signal average. Images were acquired
in the coronal, transverse and sagittal planes. The
specimen was placed in a dedicated (human) knee
coil.

X-rays were taken by Dr Ken Aplin of the
Western Australian Museum, using standard x-ray
film and processing techniques. The fine details of
the images were verified by dissection.

RESULTS AND DISCUSSION

Computed tomography was most useful for
examining hard, bony structures of the body (Figure
1). Such 3-D images of the skeleton of N. caurinus
are useful because no fully-articulated specimen of
this species is locally available, and because the
functional form of its skeleton cannot necessarily be
inferred from simple reconstruction of
disarticulated bones because of its unique fossorial
lifestyle among marsupials. Thus, 3-D computer
reconstructions of CT scans provide invaluable
information concerning the normal in situ skeletal
arrangement. X-rays were necessary to provide fine
detail of skeletal structures. MRI was useful in
viewing the internal arrangement of soft tissue
structures.

3-D Skeletal Reconstruction

The axial skeleton, comprising the skull, vertebral
column, sternum and ribs, and tail of the north-
western marsupial mole is highly adapted for its
fossorial lifestyle (Figure 1). From the lateral view
of the skeletal image, the skull is conical in shape
(Figure 1C). The skull of N. caurinus appears to be
quite thin-walled dorsally and anteriorly, but strong

in the basicranial region. Teeth are not apparent on
the CT scans because they are too small to be
resolved by a scan of 1.0 mm slice size. The skull,
including the major contours and teeth, is finely
imaged by X-ray (Figure 2). The rostrum curves
ventrally slightly, anterior to the molar teeth. There
is little evidence of the orbit or zygomatic arch. The
posterior region of the skull is smoothly curved
with no sagittal or occipital crests.

The vertebral column is remarkably strengthened,
and in lateral view has an unusually flat shape
(Figure 1C). Overall, the form of the vertebral
column appears highly specialised for burrowing.
While CT resolution is not quite adequate for the
fine detail of vertebrae, it is apparent that they are
very robust. The cervical vertebrae are obscured by
the pectoral girdle, but appear to be greatly
compressed and follow a deeply concave path to
the base of the skull. This unusual form of the
cervical vertebrae is best seen from the lateral view
of the skeletal images (Figures 1C, 2B). Fusion of
the cervical vertebrae is extremely unusual for
mammals, and is unique among marsupials for the
marsupial mole. The thoracic vertebrae are not seen
as separate entities in the 3-D skeletal images. The
shape of the vertebral column in the thoracic region
follows a sharp incline behind the pectoral girdle
and then flattens out considerably. The vertebral
column of more generalised quadrupeds follows a
wider curve than that apparent for N. caurinus
(Slijper 1946).

The lumbar region is best seen from dorsal view
(Figures 1A, 2A). The lumbar vertebrae are distinct,
becoming large posteriorly towards the pelvis.
Their transverse processes can just be discerned on
CT images. Like the posterior thoracic vertebrae, the
lumbar vertebral region is very flat in shape. The
sacral vertebrae are greatly expanded in size and
are fused with the pelvis. The wide metapophyses
are broadened to form a shield-shaped arrangement
when viewed from the dorsal aspect (Stirling, 1891;
Chapman, 1919).

An unusual feature of the vertebral column from
the dorsal view is the arrangement of vertebrae in
the tail (Figure 1). Particularly in the middle region
of the tail, these caudal vertebrae are greatly
developed; large transverse processes and chevron
bones (apparently fused to the vertebral bodies) are
evident on the CT scans. From the lateral aspect,
the caudal vertebrae are almost as large as the
thoracic and lumbar vertebrae, which is most
unusual. Generally, caudal vertebrae of mammals
are smaller and simpler in form than the thoracic or
lumbar vertebrae. The large size of the marsupial
mole's caudal vertebrae, the development of
transverse processes for muscle attachment, and the
large size of the chevron bones, suggest that the tail
is highly mobile, muscular and probably involved
in burrowing locomotion (see Withers et al., 2000).

3-D anatomy of the North-Western Marsupial Mole

3

Figure 1 Computed tomography images of the three-dimensional anatomy of the North-Western Marsupial Mole,
Notoryctes caurinus ; a dorsal, b ventral, and c lateral views. Caudal vertebrae (CaV), cranium (Cr), femur
(Fern), humerus (Hm), mandible (Ma), manubrium stemi (MS), patella (Pat), pelvis (Pel), ribs (r), scapula
(Sc), sternum (St), stomach (St), tibia (Tib), vertebral column (VC).

4

N. Warburton, C. Wood, C. Lloyd, S. Song, P.C. Withers

Figure 2 X-ray of the North-Western Marsupial Mole, Notoryctes canrinus (WAM M41482); lateral view. Actual body
length (snout-tail) = 102 mm. Legend: caudal vertebrae (Cau), chevron bones (Chev), clavicle (Clav), cranium
(Cran), femur (Fern), manus (Man), patella (Pat), pelvis (Pel), pes (Pes), radius (Rad), scapula (Sea), tibia
(Tib).

The ribs and sternum can be clearly seen in the
CT skeletal images. The rib cage is in the form of a
wide cone, being very narrow anteriorly and
expanding quite widely posteriorly (Figure 1A, B).
The ribs appear to become less dense posteriorly.
The costal cartilages, which join the ribs to the
sternum, cannot be seen. The anterior part of the
sternum is characterised by two large lateral
processes of the manubrium.

The appendicular skeleton includes the pectoral
and pelvic limbs and their girdles. The highly
modified morphology of these skeletal components
in the marsupial mole can be seen in the 3-D
skeletal reconstruction (Figure 1). Most noticeably,
the pectoral girdle is quite anteriorly placed. The
position of the shoulder articulation, of the glenoid
region of the scapula and the humerus, appears
level with the anterior cervical vertebrae, just
behind the skull, rather than the more common
position of level with the anterior thoracic vertebrae
(Hildebrand, 1988). In the lateral view, it is
apparent that the shoulder articulation is placed
quite low on the side of the body. This antero-
ventral position of the pectoral girdle is
characteristic of the position for other small
fossorial mammals (Gambaryan and Keilan-
Jaworowska, 1997).

The individual components of the pectoral girdle
can be seen with reasonable clarity (Figure 1). It is
apparent that the scapula is substantially modified,
being narrow for the most part and fanning out
posteriorly. The orientation of the scapula is also
modified from the usual dorsal arrangement of the
scapula in mammals (Hildebrand, 1988), occupying

a more laterally-facing position (Figure 1C). The
robust humerus can be seen most clearly in the
dorsal views of the skeletal image; however this
may include parts of the ulna included with the
distal end of the humerus. Unfortunately the
antebrachium (lower arm) and the manus were not
represented in the skeletal images. This is quite
unexpected as the bones of this region are relatively
more robust than, for instance, the ribs, which were
seen clearly in the skeletal reconstruction images.

The pelvic girdle is perhaps the most clear
component of the skeleton. The bones of the pelvis
are obviously highly derived and are fused to a
number of the sacral vertebrae. The typically (for
mammals) elongate innominate bones are broad
and fused with expanded elements of the sacral
vertebrae (Hildebrand, 1988). The fusion between
the pelvis and the sacral vertebrae of marsupial
moles is more extreme than in other placental
burrowers, such as armadillos and pocket gophers
(Chapman, 1919). It is evident that the pubic
symphysis is either unfused or formed by very thin
bone. The epipubic bones of marsupial moles are
vestigial and are not seen in the CT scan images.

The hind limbs of N. caurinus extend almost
laterally from the body, rather than ventrally, and
the tibiae lie flat against the ground. Mammals
generally have a parasaggital arrangement of the
limbs under the body (Hildebrand, 1988); the
abducted posture of the hindlimbs in marsupial
moles is reminiscent of a more reptilian stance. The
long bones of the limb are relatively robust, but no
fusion between the elements has taken place. An
unusual feature is the large triangular keel of the

3-D anatomy of the North-Western Marsupial Mole

5

Figure 3 Magnetic resonance image of the three-dimensional anatomy of the North-Western Marsupial Mole,
Notoryctes caurinus; sections through coronal view, dorsal to ventral. Organs visible: anal gland (A), brain
(B), fat (F), femur (Fe) heart (H), hip musculature (Hp), humerus (Hu), intestine (I) liver (Li), lungs (Lu), nose
shield (N), oral cavity (O), rib cage (R), shoulder musculature (Sh), stomach (St), tail musculature (Ta), testes
(T), thigh musculature (Th), and vertebral column (V).

patella at the knee joint of the hind limbs (Figure
1C). Few other marsupials (only bandicoots) have
an ossified patella, and none have one resembling
the relatively large, irregular patella of Notoryctes.
The tibia and fibula are pictured for the right leg
only, but are clearly quite robust for an mammal of
this size. Unfortunately, like the forelimb, the distal

parts of the hind limb are not pictured in the 3-D
skeletal image. Again this is unexpected as they
have relatively robust bones. The unusual
morphology of the hind limb, with the large
triangular patella and expanded tibia is more
clearly visible in the X-ray than the CT scan images.
Like the carpus of the forelimb, the small bones of

6

the pes are highly modified and difficult to identify
even from X-ray images. However, it is interesting
to note that the clawed digits of the foot are
splayed, and held in a somewhat abducted and
medially rotated posture, presumably to achieve the
maximum surface area of the sole of the foot when
burrowing.

Soft Tissue Structure

From all MRI views (Figure 3), the most apparent
soft-structure is a large amount of subcutaneous fat
on this specimen, particularly around the ventral
surface of the pelvis but also dorsal to the pelvis
and anteriorly around the shoulders. This very large
amount of fat appears as a high signal intensity on
the magnetic resonance images. Upon dissection,
there was approximately four grams of
subcutaneous fat - more than 10% of a 35 gram
animal.

The largest organs of the digestive system, the
liver, stomach and intestine, are easily
distinguished. The abdominal cavity appears as
very low signal intensity, oval-shaped structure in
the mid-region of the animal, due to this space
being air-filled as a result of an incision made in the
skin for the penetration of formalin /alcohol for
fixation of the internal organs. The liver is the most
anterior organ of the abdomen, and multi-lobed in
structure. The stomach appears quite dark on the
left side of the abdominal cavity in the coronal
sections. Also seen from CT scans, the stomach
contains a large amount of sand resulting in the low
signal intensity in MRI. The intestine fills the rest of
the abdominal cavity and has a higher signal
intensity than the stomach and liver. The smaller
organs of the abdominal cavity, spleen, gall bladder,
etc are not distinguishable. The oral cavity is also
visible in the relevant coronal sections (Figure 3).

In the thoracic cavity, the heart is the only portion
of the circulatory system that is clearly apparent in
MRI. At particular levels of contrast it is possible to
discern the chambers of the heart. Of the respiratory
system the lungs are visible. Their triangular shape is
best observed from the coronal sections (Figure 3).

The urinogenital system is poorly represented in
the MRI due to the small size of these organs; only
the testes are clear (in all views) and the kidneys in
sagittal view. Posteriorly, lying in the mass of fat
under the pelvis, the testes are seen very clearly,
with good contrast between gonadal tissue and fat.
It is difficult to assess the sex of marsupial moles
externally as males have abdominal testes, so MRI
provides a non-intrusive method for confirming
that this specimen is male.

The brain is the only obvious component of the
nervous system in the MRI images (Figure 3). With
the MRI having a slice size of 1.4 mm, it is possible
that the spinal cord was missed altogether in
coronal and sagittal sections.

N. Warburton, C. Wood, C. Lloyd, S. Song, P.C. Withers

The hard, bony structures of the skeletal system
appear dark on the MRI. Clearly visible are sections
through the skull, vertebral column, ribs, humeri
and femora. The large size of the vertebrae is
obvious and it is easier to distinguish adjacent
vertebrae in MRI than CT scans. Surrounding the
long bones of the limbs, the major muscle groups
are visible, but distinction between individual
muscles is not possible except for the very large
muscles of the thigh, upper arm and base of the tail.
In these areas the muscle bulk is enormous, much
greater than for non-fossorial mammals of
comparable size ( personal observations). The
extensive muscles attaching to the tail are quite
unusual and would seem to be related to the
strengthening of the caudal vertebrae as noted from
the CT skeletal images. The muscles of the tail also
appear to be more strongly developed ventrally (for
flexion) than dorsally (for extension).

In summary, traditional X-rays provide detail of
skeletal elements but are sometimes difficult to
interpret because a three dimensional structure has
been projected onto a two dimensional image.
Reconstruction of CT scans results in a three
dimensional image that reduces the problem of
overlap. Also, with the CT skeletal reconstruction, it
is possible to rotate the image to obtain the desired
view, an impossible manipulation with X-ray
images. MRI provides information of the anatomy
of soft tissue structures by recording the differential
contrast of the various soft tissues. Due to the small
size of the marsupial mole, detail in sections was at
the resolution of the MR scanner. Nevertheless,
some valuable information concerning the 3-D
arrangement of the soft anatomy is apparent.

ACKNOWLEDGEMENTS

We are grateful to Ms Norah Cooper and Dr Ken
Aplin of the Western Australian Museum for the
loan of the specimen, and the X-ray images.

REFERENCES

Chapman, R. N. (1919). A study of the correlation of the
pelvic structure and the habits of certain burrowing
mammals. American Journal of Anatomy 25: 185-219.
Gambaryan, P. and Kielan-Jaworowska, Z. (1997).
Sprawling versus parasagittal stance in
multituberculate mammals. Acta Palaeontologica
Polonica 42(1): 13-44.

Hildebrand, M. (1988). Analysis of vertebrate structure
(third edition). John Wiley and Sons, Inc. New York.
Rogers, S. W. (1999). AUosaurus, crocodiles and birds:
evolutionary clues from spiral computed tomography
of an endocast. Anatomical Record 257(5): 162-173.
Romagnano, A., Shiroma, J. T., Heard, D. )., Johnson, R.
D., Sheiring, M. R., and Mladinich, C. (1996).
Magnetic resonance imaging of the brain and
coelomic cavity of the domestic pigeon ( Columba livia

3-D anatomy of the North-Western Marsupial Mole

domestica). Veterinary Radiology and Ultrasonnd 37(6):
431-140.

Samii, V. F v Biller, D. S., and Koblik, P. D. (1999).
Magnetic resonance imaging of the normal feline
abdomen: an anatomic reference. Veterinary Radiology
and Ultrasound 40(5): 486—190.

Slijper, E.J. (1946). Comparative biologic-anatomical
investigations on the vertebral column and spinal
musculature of mammals. Koninklijke Nederlandsche
Akademie van Wetenschappen., Verhandelingen
(Tweede Sectie) 42: 1-128.

Stirling, E. C. (1891). Description of a new genus and
species of Marsupialia, Notoryctes typhlops.
Transactions of the Royal Society of South Australia 14:
154-187.

7

Thomas, O. (1920). Notoryctes in north-west Australia.

Annals and Magazine of Natural History (9) 6: 111-113.
Thompson, G. G., Withers, P. C. and Seymour, R. S.
(2000-2001). Blind diggers in the desert. Nature
Australia (Summer) 26-31.

Withers, P. C., Thompson, G. G. and Seymour, R. S.
(2000). Metabolic physiology of the north-western
marsupial mole, Notoryctes caurinus (Marsupialia:
Notoryctidae). Australian Journal of Zoology 48: 241-
258.

Manuscript received 6 November 2002; accepted 9 June 2003

Records of the Western Australian Museum 22: 9-16 (2003).

Description of Volutoconus hargreavesi calcarelliformis subsp. nov.
(Mollusca: Volutidae) from northwestern Australia

Fred E.Wells and Hugh Morrison

Department of Aquatic Zoology
Western Australian Museum
Perth, Western Australia 6000, Australia

Abstract - Volutoconus hargreavesi calcarelliformis subsp. nov. is described from
the outer continental shelf off Port Hedland, northwestern Australia. The new
subspecies is compared with V. hargreavesi hargreavesi (Angas, 1872), and V.
hargreavesi daisyae Weaver, 1967, which is recognised as a valid subspecies.

Keywords: Subspecies, volute, Volutoconus, Gastropoda, Volutidae, Western
Australia

INTRODUCTION

The volute genus Volutoconus is a small group of
five species which are restricted to northern
Australia, adjacent areas of the east and west coasts
of the continent, and eastern Indonesia. The type
species, V. coniformis (Cox, 1871), occurs along the
north coast of Western Australia from the Dampier
Archipelago to the west Kimberley, Further north,
V. bednalli (Brazier, 1878) extends across the north
Kimberley to Torres Strait, Queensland and
throughout the Timor Sea to eastern Indonesia,
including Irian Jaya. Volutoconus grossi (Iredale,
1927) ranges southward along the east coast of
Australia from Townsville, Queensland to Port
Macquarie, New South Wales. Two subspecies are
generally recognised: V. grossi grossi (Iredale, 1927),
which occurs south from Keppel Bay, Queensland
and V. grossi mcmichaeli Habe and Kosuge, 1966,
which occurs further north in the Townsville area.
The fourth species, V. hargreavesi Angas, 1872,
occurs widely along the west coast of Western
Australia, from Mandurah to North West Cape, and
along the north coast as far as the western
Kimberley. Another species, V, capricomeus, was
described by Wilson (1972) from west of Point
Cloates and provisionally placed in Volutoconus.
The generic placement of the species has varied, but
the most recent analysis (Willan, 1995) places it in
Volutoconus.

Weaver (1967) described a geographical variant
of Volutoconus hargreavesi as the subspecies V.
hargreavesi daisyae, which occurs along the west
coast of Western Australia from North West Cape
south to about Mandurah. Weaver restricted the
nominate subspecies, V. hargreavesi hargreavesi, to
the north coast of the state. However, Weaver and
DuPont (1970) subsequently reported that further

collecting had narrowed the gap between the two
forms, and that subspecies status was not justified.
Wilson and Gillett (1971) retained the separation of
the subspecies, but stated that the finding of
intermediates might show that the separation was
not warranted. Wilson (1994) later considered that
there was no need to separate the two subspecies.
Poppe and Goto (1992) separated the two, but as
forms, which have no taxonomic standing.

Recent curation of the volutes in the collection of
the Western Australian Museum has uncovered a
deep-water subspecies in the V. hargreavesi
complex, which is described here. In addition, V.
hargreavesi daisyae is recognised as a valid
subspecies.

Institutional acronyms: AMNH, American
Museum of Natural History; DMNH, Delaware
Museum of Natural History; NTM, Northern
Territory Museum, WAM, Western Australian
Museum.

SYSTEMATICS
Family Volutidae
Genus Volutoconus Crosse, 1871

Volutoconus Crosse, 1871: 306

Type species

Voluta coniformis Crosse, 1871, by original
designation.

Diagnosis

(Adapted from Weaver and DuPont [1970]).
Moderately large (60 to 130 mm) volutes, solid, with
a range of colour patterns. Spire variable, elevated

10

F.E. Wells, H. Morrison

Figure 1

Distribution of Volutoconus Iwrgreavesi calcarelliformis subsp
1/. hargreavesi daisyae Weaver, 1967.

. nov., V. hargreavesi hargreavesi (Angas, 1872), and

Volutoconus hargreavesi from Western Australia

11

Plate 1 Holotype of Volutoconus hargreavesi calcarelliformis subsp. nov. (WAM S 14359)

or depressed. Protoconch smooth or radially ribbed,
usually possessing a sharp apical spire (calcarella).
Columella with four or more plaits. Periostracum
and operculum absent. Radula (where known)
uniserial, with large, tricuspid teeth, cusps strongly
arched and fang like, median cusp much longer
than laterals.

Remarks

The genus is restricted to northern Australia and
Indonesia, ranging over the entire northern
coastline of the continent, with individual species
extending down both the west and east coasts. The
systematics of Volutoconus have been examined in
detail by Abbott (1958); McMichael (1960); Weaver
(1967); Weaver and duPont (1970); Wilson and
Gillett (1971); Wilson (1972; 1994) and Poppe and
Goto (1992). The genus occurs from intertidal areas
to at least 200 m. Fossils previously included in the
genus were considered to belong to Nannamoria by
Darragh (1988). Darragh recorded only a single
fossil specimen of an unidentified Volutoconus,
which he considered to closely resemble V.
hargreavesi.

Volutoconus hargreavesi hargreavesi (Angas, 1872)
Plates 2c, 3c, Figure 1

Voluta hargreavesi Angas, 1872: 613-614, plate 42,
figure 13 (locality unknown); Weaver, 1960: 1, 3
front page plate (4 shells on left); Weaver, 1967:
302, 304, plate 41, figures 1-4, map; Weaver and
duPont, 1970: 138, plate 61E-F; Matsukuma et
al, 1991: p. 99, plate 97, figure 3; Poppe and
Goto, 1992: 212, plate 107, figures 3-5, 7.

Cymbiola ( Volutoconus ) hargreavesi (Angas): Abbott,

' 1958:4

Volutoconus hargreavesi hargreavesi (Angas): Wilson
and Gillett, 1971: 74, plate 49, figure 5.

Type material

Location of type unknown.

Weaver (1967) states the holotype is lodged in the
American Museum of Natural History, number
AMNH 8304. Weaver & DuPont (1970) questioned
whether this is in fact the holotype as there are
differences between the colour pattern on the shell
and the illustration.

The published catalogue of the AMNH (Boyko and

12

F.E. Wells, H. Morrison

Comparison of shells of Volutoconus hargreavesi. a. Holotype of V. hargreavesi calcarelliformis subsp. nov.
(WAM S 14359). b. V. hargreavesi daisyae Weaver, 1967 (WAM S 14358). c. V. hargreavesi hargreavesi (Angas
1872) (WAM S 14348).

Cordeiro, 2001) states: "AMNH 8304. ex Steward
Collection #1971. Richards and Old (1969: 117)
summarized the limited information about this shell.
They noted that it did not match exactly the size
given by Angas (1872) for his unique specimen.
AMNH 8304 is the same size as the figure of the
shell in Angas (1872: plate 42, figure 13) and matches
the description. However, the AMNH shell is much
more worn than suggested by Angas's (1872) figure,
and the color pattern is quite different. Based on the
available evidence, we are inclined to consider
AMNH 8304 an old specimen of this taxon, but we
think its holotype status is unlikely."

Type locality

Angas (1872) did not provide a type locality for

V. hargreavesi. Weaver (1960) subsequently
designated Bedout Island in the Dampier
Archipelago as the type locality.

Material examined: Australia: Western
Australia: WAM S 14348, 2 specimens, NW of
Roebuck Bay, S of Lacepedes Island (16°52'S;
122°08'E); WAM S 14349, 1 specimen, off Cape
Lambert (20°36'S; 117°10'E); WAM S 14350, 1
specimen, 26 m, Dampier Archipelago,
approximately 18 km W of Eaglehawk Island,
(20°39'S;116°18’E); WAM S 14351, 1 specimen, 97
m, Onslow, 13 km N of Long Island
(20 a 55'S;115°51'E); WAM S 14352, 1 specimen, no
data (seized by Australian Customs); WAM S
12669, 1 specimen, 103 m. North West Cape
(21°36.22'S; 114°11.11'E), on sand and mud.

Volutoconus hargreavesi from Western Australia
Diagnosis

Calcarella small, orange colour, very finely
ribbed. First two whorls of teleoconch white with
very fine axial ribbing. Upper whorls with very fine
ribbing. Body whorl with distinct, but fine ribbing.
Colour variable, mottled orange and white,
generally with 2 indistinct bands, one on shoulder
of body whorl, one at base of whorl, sometimes a
third intermediate band is present at base of body
whorl. Four high, narrow columellar plaits.
Anterior canal with deep notch, margin raised.
Interior pale fawn to white.

Range

North West Cape to west Kimberley, Western
Australia

Depth range

Intertidal to about 100 m.

Animal

Not known.

Remarks

The three subspecies of V. hargreavesi are
geographically distinct (Table 1). The nominate
subspecies, V. hargreavesi hargreavesi, occurs along
the continental coastline, from about North West
Cape to the west Kimberley. Most specimens are
from shallow water, but there are two lots in the
WAM from about 100 m. Volutoconus hargreavesi
daisyae occurs further to the south, along the west
coast of Western Australia, from about Mandurah
to North West Cape, in depths of 40 to 134 m. There
is an area at North West Cape where the species
ranges may or may not overlap. In contrast, V.
hargreavesi calcarelliformis lives in deeper waters

13

(150-202 m) on the North West Shelf from the
Rowley Shoals to Cartier Island.

Compared to V. hargreavesi calcarelliformis, V.
hargreavesi hargreavesi has a smaller calcarella for the
size of the shell. The shell of V. hargreavesi
hargreavesi is bigger with a smoother surface.
Volutoconus hargreavesi hargreavesi differs from V.
hargreavesi daisyae in being a broader shell with fine
growth lines instead of the more pronounced ribs
found on V. hargreavesi daisyae.

Volutoconus hargreavesi daisyae Weaver, 1967
Plates 2b, 3b

Volutoconus "species" Angas, Weaver, 1960: 1, 3
front page plate (large centre shell).

Volutoconus hargreavesi daisyae Weaver, 1967: 302,
304, plate 41, figures 5-9, map; Wilson and
Gillett, 1971: 74, plate 49, figure 6.

Volutoconus hargreavesi (Angas): Weaver and
duPont, 1970: 138, plate 61G; Poppe and Goto,
1992: 212, plates 106, figure 1,2; Wells and Bryce,
1986: 118, figure 456; Wilson, 1994: 118, plate 21,
figure 4 a-e.

Type material

Holotype, DMNH 10022.

Type locality

Southwest of North West Cape, central Western
Australia, 22°00'S; 113°45'E.

Material Examined

Australia: Western Australia: WAM S 14355, 1
specimen, 134 m, NW of Shark Bay (23°39'S;
113°11'E); WAM S 14356, 1 specimen, 134 m, NW of

Plate 3 Comparison of protoconchs of Volutoconus hargreavesi. a. Holotype of V. hargreavesi calcarelliformis subsp. nov.

(WAM S 14359). b. V. hargreavesi daisyae Weaver, 1967 (WAM S 14358). c. V. hargreavesi hargreavesi (Angas,
1872) (WAM S 14348).

14

F.E. Wells, H. Morrison

Shark Bay (23°39'S; 113°11'E); WAM S 14357, 1
specimen, 110 m, 92 km W of Dongara (29°07.5'S;
113°57.4'E), sponges and stone rubble; WAM S
14358, 1 specimen, off Greenhead (30°04'S
114°58'E).

Diagnosis

Calcarella small, orange colour. Shell fusiform.
First two whorls of teleoconch white with strong
axial ribbing. Upper whorls with very strong
ribbing extending onto body whorl. Body whorl
heavily ribbed. Colour variable, mottled orange and
white with 2 indistinct bands, one on upper body
whorl, one lower on whorl. Four high, narrow
columellar plaits. Anterior canal with deep notch,
margin raised. Interior pale fawn to white.

Range

West coast of WA; Mandurah to North West
Cape, Western Australia

Depth

Approximately 40 m to 134 m.

Animal

Not known.

Remarks

Volutoconus hargreavesi daisyae differs from V.
hargreavesi hargreavesi in having a narrower shell
with pronounced axial ribbing instead of the fine
growth lines found on V. hargreavesi hargreavesi.
Volutoconus hargreavesi daisyae differs from V.
hargreavesi calcarelliformis in being longer, more
robust, with a similar protoconch, but a brighter
orange colour. The ribs of V. hargreavesi daisyae are
much heavier.

Volutoconus hargreavesi calcarelliformis
subsp. nov.

Plates 1, 2a and 3a

Holotype

WAM S 14359, 150-160 m, 239 km NE of Cape
Lambert (18°42.8'S; 118°03.2'E), Western Australia,
Australia, sand, rubble and dead shells, collected
by L.M. Marsh et. al. on 18 August 1995.

Paratypes

Australia: Western Australia: WAM S 14360, 6
specimens, 173-193 m, 70 km S of Cunningham
Island, Imperieuse Reef, Rowley Shoals (18°06.9'S;
118°56.7'E), sand, rubble and dead shells; WAM S
14361, 5 specimens, 150-160 m, 239 km NE of Cape
Lambert (18°42.8'S; 118°03.2’E), rubble and dead
shells; WAM S 14362, 1 specimen, 150-160 m, 239
km NE of Cape Lambert (18°38.6'S; 118°07'E), sand.

rubble and dead shells; WAM S 14363, 2 specimens,
154 m, 184 km N of Port Hedland (18°47'S;
117°58'E), grey mud and shell rubble; WAM S
14364, 2 specimens, 154 m, 184 km N of Port
Hedland (18°47'S;117°58'E), grey mud and shell
rubble; WAM S 14365, 1 specimen, 201-202 m, 217
km N of Port Hedland (18°20.5'S; 118°27.7'E), grey
mud; WAM S 14389, 1 specimen, 150-160 m, 239
km NE of Cape Lambert (18°42.8'S 18°39.6'S), sand
rubble and dead shells. NTM P8581, 1 specimen,
180 m, 20 km south of Barracouta Shoal, west of
Cartier Reef, Sahul Shelf (12°42.86'S; 123°57.98’E),
coarse broken shell substrate; Mike Claydon private
collection, 1 specimen, trawled by scampi boats, on
sand mud and shell, 220 m, NW of Point Samson.

Description

Calcarella prominent, tall, smooth sided, lacking
axial ribbing. Bright orange colour. First two whorls
of teleoconch white, with strong axial ribbing.
Lower whorls also heavily ribbed with mottled
fawn to orange patterning. Body whorl finely
ribbed, mottled brown with 3 faint bands, one
below suture, one on outer body whorl, one lower
on whorl. Four high, narrow columellar plaits.
Anterior canal with deep notch. Interior white,
some shells with pale fawn to white on inside of
outer lip.

Shell measurements

Table 1 shows the measurements of the type
series.

Geographical range

North West Shelf, Western Australia, from about
Rowley Shoals to Cartier Island.

Depth

150-202 m.

Animal

Not known.

Etymology

Named after the distinctive calcarella at the apex
of the shell spire.

Remarks

The key feature of Volutoconus hargreavesi
calcarelliformis is the prominent size of the calcarella
for the size of the shell. It differs from V. hargreavesi
hargreavesi in being smaller, with fine ribbing,
strongly ribbed protoconch, and it lives in slightly
deeper water. Volutoconus hargreavesi hargreavesi has
a smoother shell with fine growth lines. Volutoconus
hargreavesi calcarelliformis differs from V. hargreavesi
daisyae in being shorter, more robust, with a similar
protoconch, but a brighter orange colour. The ribs

Volutocomis hargreavesi from Western Australia

15

16

F.E. Wells, H. Morrison

of V. hargreavesi daisyae are much stronger than
those of V. hargreavesi calcar elliformis.

The dwarf form of V. grossi (Iredale, 1927) from
deep water off the Great Barrier Reef, Queensland,
is very similar to V. hargreavesi calcarelliformis.
Further material is required before the relationships
between V. grossi and V. hargreavesi can be fully
understood. In addition to the considerable
geographic separation with a shallow water barrier
between them, dwarf specimens of V. grossi can be
separated from V. hargreavesi calcarelliformis by
having a narrower shell and finer ribbing, and
white calcarella.

ACKNOWLEDGEMENTS

We are grateful to Dr Richard Willan for
information on the new subspecies, and Mike
Claydon for the loan of material of the new
subspecies. Clay Bryce was kind enough to
photograph the specimens. We are pleased to
acknowledge discussions with Dr Patrice Bail, Glad
Hansen, Alan Limpus, and Dr Barry Wilson. Corey
Whisson databased the specimens and assisted with
preparation of the figure and plates. Drs Tom
Darragh and Richard Willan very kindly reviewed
the maunscript.

REFERENCES

Abbott, R.T. (1958). Notes on the anatomy of the
Australian volutes, bednalli and grossi. Journal of the
Malacological Society of Australia 1(2): 2-7.

Angas, G.F. (1872). Description of a new species of
Voluta. Proceedings of the Zoological Society of London
1872: 613-614.

Boyko, C.B. and Cordeiro, J. (2001). Catalog of Recent
Type Specimens in the Division of Invertebrate
Zoology, American Museum of Natural History. V.
Mollusca Part 2 (Class Gastropoda [exclusive of
Opisthobranchia and Pulmonata], with Supplements
to Gastropoda [Opisthobranchia], and Bivalvia).
Bulletin of tire American Museum of Natural History 262:
1-170.

Brazier, John. (1878). Descriptions of seven new species
of terrestrial and marine shells from Australia.
Proceedings of the Zoological Society of London 1878: 20-
23.

Cox, J.C. (1871). Description de deux especes novelles de
Voluta et observations sur le V. punctata Swainson.
Journal de Conchyliologie 3(19): 74-77.

Crosse, J.C.H. (1871). Distribution geographique et
catalogue des especes actuelles du genre Voluta.
Journal de Conchyliologie 14: 105-117.

Darragh, T.A. (1988). A revision of the Tertiary Volutidae
(Mollusca: Gastropoda) of south-eastern Australia.
Memoirs of the Museum of Victoria 49: 195-307.

Iredale, T. (1927). Caloundra shells. The Australian
Zoologist 4: 336.

McMichael, D.F. (1960). Notes on some Australian
Volutidae. Journal of the Malacological Society of
Australia 1(4): 4-13.

Matsukuma, A., Okutani, T., and Habe, T. (1991). World
Seashells of Rarity and Beauty. National Science
Museum, Tokyo.

Poppe, G.T. and Goto, Y. (1992). Volutes. Mostra
Mondiale Malacologia, Cupra Marittima (AP- Italy).
Richards, M.C. and Old, W.E. Jr. (1969). A catalogue of
molluscan type specimens in the Department of Living
Invertebrates, the American Museum of Natural History.
American Museum of Natural History, New York.
Unpublished report, 147 pp.

Weaver, C.S. (1960). Hawaiian scientific expedition finds
rare Australian volutes. Hawaiian Shell News 8(10): 1,3.
Weaver, C.S. (1967). A new subspecies of Volutoconus
hargreavesi (Angas, 1872) from central Western
Australia. The Veliger 9(3): 301-304.

Weaver, C.S. and duPont, J.E. (1970). The Living Volutes.
A monograph of the Recent Volutidae of the world.
Delaware Museum of Natural History, Monograph
Series No. 1: 1-375.

Wells, F.E. and Bryce, C.W. (1986). Seashells of Western
Australia. Western Australian Museum, Perth.

Willan, R.C. (1995). Taxonomic and biogeographic review
of the Australian endemic genus Nannamoria
(Gastropoda: Volutidae) with the description of a new
bathyal species. Invertebrate Taxonomy 9: 107-113.
Wilson, B.R. (1972). New species and records of
Volutidae (Gastropoda) from Western Australia.
Journal of the Malacological Society of Australia 2(3): 339-
360.

Wilson, B.R. (1994). Australian Marine Shells. Volume 2.

Odyssey Publishing, Kallaroo, Western Australia.
Wilson, B.R. and Gillett, K. (1972). Australian Shells. A.H.
and A.W. Reed, Sydney.

Manuscript received 29 April 2003; accepted 29 July 2003

Records of the Western Australian Museum 22: 17-22 (2003).

A new genus and species of trigoniuline milliped from Western Australia
(Spirobolida: Pachybolidae: Trigoniulinae)

Richard L. Hoffman

Virginia Museum of Natural History
Martinsville, Virginia 24112, USA

Abstract - A new genus and species, Austrostrophus stictopygus, is described
from northern Western Australia. The presence of punctate paraprocts, very
rare in spiroboloid diplopods, is shared with Ainigmabolus chisholmi of New
South Wales, but apparently only as a homoplasic condition.

INTRODUCTION

Whether considered a subfamily of the
Pachybolidae or a discrete family, trigoniuline
millipeds comprise a conspicuous element in the
fauna of southeastern Asia and Indonesia, whence
some 21 genera are currently recognized (Jeekel,
2001). The group is, however, only marginally
represented in Australia, chiefly by Zygostrophus
and Ainigmabolus on the eastern edge of the
continent, and the recently described Speleostrophus
(Hoffman, 1994) from an island on the west coast.
The recent discovery of a highly disjunct new
trigoniuline from Western Australia suggests that
undercollecting along the western and northern
periphery of the continent may account for this
apparent poverty of Australian spiroboloids.

Family Pachybolidae Cook
Subfamily Trigoniulinae Attems
Austrostrophus gen nov

Type species

Austrostrophus stictopygus sp. nov.

Diagnosis

A trigoniuline genus characterized by the
enlarged, distally excavate, gonopod coxae, into
which the elongated, slender telopodites are basally
recurved. Submarginal belt of paraprocts
conspicuously set with large coarse punctures.
Posterior edge of metazona with specialized fringe
composed of small denticulate projections.

Distribution

Western Australia.

Etymology

A masculine neologism composed of the elements

austro (from Latin australis, in allusion to the
country of origin) + -strophus, a combining form
employed in many generic names in this subfamily.

Austrostrophus stictopygus sp. nov.

Figures 1-10

Material examined

Male holotype (WAM T46241), from Western
Australia: Burrup Peninsula, Rocky Hill at Hearson
Cove (20°37'06"S, 116°47'05"E), R. Teale leg. 22 May
2002. Two immature females (WAM T46956) from
the same locality, 30 September 2002, R. Teale and
M. Maier, leg.

Diagnosis

With the characters of the genus.

Holotype

Adult male, length approximately 52 mm
(broken), maximum diameter 4.2 mm. 51 segments
+ telson. Colour of recently preserved specimen:
prozona grayish, mesozona piceous to black,
metazona orange-brown. Mesozonal black pattern
narrowed ventrad and absent above leg bases;
antennae, legs, sterna and pleural region similar in
colour to metazona.

Head without modifications, sides of genae
slightly convex but convergent ventrad;
interantennal and interocellarial widths 1.2 mm,
ocellaria subrounded, maximum length 0.5 mm,
about 36 ocelli in rows (4-5-7-6-6-5-3). 7+7 labral
setae, 2+2 clypeal setae, those of each side widely
separated. Antennae short, 2 nd and 6 th articles
longest, similar in size and shape, 3rd-5th shorter
and slightly more triangular in outline; outer distal
ends of 5 th and 6 th articles with rounded, slightly
convex sensory area, depigmented and appearing
semimembranous; 7 th article very short, with four
terminal sensory cones. Mentum of

18

R.L. Hoffman

Figure 1

Posterior end of body, showing abbreviated form of the epiproct and coarsely punctate condition of the
paraprocts (SEM x32).

Figure 2 Posterior edge of metazonum showing the palaeate fringe (SEM x200).

gnathochilarium with low but distinct median
ridge, prebasilar sclerite very thin. Mandibular
bases without lobes or other modification.

Lateral ends of collum subtriangular as usual in
this family; pleural region of 2 nd segment produced
ventrad and visible beneath end of collum as a
digitiform lobe. Surface of collum and all body
segments superficially smooth and polished: with
magnification (60X) surface texture is a uniform
isodiameteric mesh, on which is superimposed a

pattern of longitudinal striations; prozona with
several concentric striations below level of
ozopores, curved posteriad at their ventral ends.
Pores prominent, located in mesozona below level
of lateral suture. Pleurotergal suture line visible;
sterna subquadrate, smooth (Figure 4). Metazona
slightly elevated above level of mesozona, and
caudally decurved slightly, becoming much thinner
and edged with a fringe composed of small
separate, denticulate units (paleae) (Figures 2, 3).

A new genus and species of trigoniuline milliped

19

3

5

Figures 3-5 3. Two individual paleae, greatly enlarged (freehand, about 2000 x). 4. Sternal region of midbody segment,

legs removed to show coxal sockets and partial closure of posterior sockets. 5. Bases of leg pairs 2-6,
showing form of processes of 3 rd and 4 lh coxae, posterior aspect.

Terminal segment (Figure 1) not produced
dorsomedially; most of paraprocts visible in dorsal
aspect, their posterior edge set off by a moderately
deep, broad groove beset with numerous large
coarse punctures, disk of paraprocts impunctate or
nearly so. Hypoproct transversely narrow,
unmodified.

Posterior coxal cavities partially closed by small
median projection of sterna (Figure 4). Legs
relatively long, tarsi visible in dorsal aspect when
extended; length ratio of podomeres in descending
order: 2-3-6-4-5-1. Podomeres essentially glabrous,
basal four with a single small seta at distal end
ventrally, postfemur with a distinctly larger seta in
that position, tarsi typically with three large setae
(nearly length of tarsal claw) in a ventrodistal row,
and a single large supra-apical seta. Anterior legs
without pads, although ventral surface of apical
podomeres appears depigmented and less
sclerotized than dorsal surface (perhaps endowed

with moderate holding ability). Tarsal claws similar
in size and shape to those of other legs. Coxae of
legs 3 and 4 produced into conspicuous, flattened
lobes, those of 4 th coxa turned laterad apically
(Figure 5).

Anterior gonopods distinctive in the globosely
enlarged coxae, deeply excavate on mesal side,
accommodating basal half of telopodites, anterior
surface of coxae not produced into slender apices
(Figure 6). Sternum modified "Y" shape, prolonged
slightly behind coxae, sternal apodeme reflexed
caudad and not visible in anterior aspect; posterior
extension of sternum separated as a discrete
elongate sclerite, narrowed at each end, and in
contact with basal end of telopodite (Figure 8, PXS).
Telopodite differing from typical trigoniuline
structure in being attached to an elevated posterior
rim of the coxa, then abruptly recurved into cavity
of the latter, distal half reflexed ventrad behind
median sternal projection as a slender stalk, apically

20

R.L. Hoffman

Figures 6-8 6. Left side of anterior gonopods, anterior aspect. S, sternum, C, coxa, T, telopodite. 7. Lateral aspect of

gonopods (posterior pair concealed) with prostatic gland (stippled, at left) and retractor muscles of
telopodite indicated by broken lines inside coxa. 8. Posterior view of left anterior gonopod, showing
prolongation of coxa (C), with base (TB) and distal shaft (TD) of telopodite. PXS, posterior extension of
sternum; SA, reflexed sternal apodeme.

capitate just beyond apex of sternal projection
(Figures 7, 9).

Posterior gonopod (Figure 10) generally
conforming to basic trigoniuline pattern: median
sternal element (S) present, merging each side into
a coalesced coxostemal region containing (1) a small
chamber into which the prostatic duct (PD)

discharges, and (2) a small, short, digitiform
projection. Distal 2/3rds of gonopod (telopodite) set
off at nearly a right angle by a deep flexible
articulation (A). Prostatic groove courses on medial
side to a clear membranous subapical lappet; form
of telopodite and its various lobes and processes as
shown in Figure 10.

A new genus and species of trigoniuline milliped

21

Figure 9 Oblique posteromedian aspect of gonopods to
show the prominent flexure of the telopodites
into the distal coxal cavity unique to this
genus.

Figure 10 Posterior gonopod, lateral aspect, sternal
apodeme concealed by the protractor and
retractor muscles. PD, prostatic duct with its
sclerotized internal tube. S, sternum; A,
flexible articulation between coxal and
telopodital regions.

Etymology

From the Greek stiktos (punctured) + pygos
(rump), in allusion to the coarsely punctate
paraprocts.

COMMENTS

Punctate paraprocts

My initial impression, confirmed by a preliminary
literature survey and survey of material at hand,
was that this character was not known for the entire
order Spirobolida. Only through serendipity did I
notice the remark by Verhoeff (1937: 149) that in
Ainigmabolus chisholmi the “ Analklappen neben dem
innenrande mit mehreren, unregelmassigen
Eindriicken." This species, a trigoniuline from New
South Wales, shows virtually no similarities in
gonopod structure with A. stictopygus, and
apparently the paraproct modification must be
considered a "geographic homoplasy" peculiar to
Australia rather than any kind of synapomorphy
for the two taxa.

Metazonal fringe

In many orders of Diplopoda the posterior edge
of the metazona is provided with a thin,
transparent, sclerotized membrane ("limbus"),
variable both in width and ornamentation of its
edge. The definitive treatment of limbus occurrence
throughout helminthomorph Diplopoda (Schmidt,
1962), recorded that in the three "trigoniuline"
families (Pachybolidae, Trigoniulidae,
Spiromimidae) "ein eigenlicher Limbus ist auch hier
nicht vorhanden" - at least in species of the six
genera examined. In one, Trigoniulus ceramicus, the
edge of the metazona itself is irregularly indented
and darkly pigmented, but the typical limbus
structure of a clear membrane with modified edge
is in no way approximated. The condition in
Austrostrophus is perhaps an elaboration of this
simple form. The margin of the metazona is notably
thinner than the sclerite otherwise, but is opaque
and pigmented, no clear membrane is present, and
the scale-like individual units (for which the term
paleae is suggested, singular palea) of the fringe
originate separately from the edge itself. Perhaps
this expression could be distinguished by the name
"paleate". In those taxa in which a true limbus is
well-developed, e.g., Odontopygidae, the hyaline
membrane itself can usually be removed intact as a
continuous clear strip. In a general way, the true
limbus seems to be best developed in taxa in which
other character systems are clearly derived, and
therefore indicative of an evolutionarily advanced
status for such groups.

Gonopods

Although it seems generally appreciated that the

22

R.L. Hoffman

sperm transfer process in most helminthomorph
diplopods is facilitated by a what is regarded as a
prostatic secretion (with ?enzymatic, ?nutritive,
?buffering function) the glands themselves have not
received a thorough comparative study. In
Polydesmida, they lie in the coelom of segment 7
(often adjacent segments as well) and their secretion
is conducted through the gonocoxae to a prostatic
groove originating at the base of the telopodite. In
Spirostreptida, the glands are generally located on
the posterior side of and in close proximity to the
gonocoxal folds. They have been scarcely noticed in
Spirobolida, but their sclerotized ducts entering the
bases of posterior gonopods are frequently
illustrated. Because of the excellent state of
preservation of the holotype of A. stictopygus I was
able to determine that the glands are located in the
8 th segment well behind the bases of the gonopods
and each is nearly as large as the preceding
gonocoxa (Figure 7). A long duct connects to the
coxosternal region, the central sclerotized tube
surrounded by a thick layer of what appears to be
additional glandular tissue (Figure 10).

The function of the small internal chamber
(Spermagrube of Verhoeff, 1937, Figure 13) remains
unknown; it appears too small to be an effective
reservoir. Some trigoniuline genera are credited
with the presence of two such chambers.

Figure 8 shows the unique formation of the
anterior gonocoxa in this genus. The posterior
surface is greatly elongated distal instead of being
short and basically transverse with the telopodite
hinged along an exposed seam and apparently
capable of very limited movement. In
Anstrostrophus, the base of die telopodite is only
narrowly joined to the edge of the coxa, and
apparently capable of being extended and retracted
(Figure 7 shows retractor muscles only, I did not
wish to locate the extensors by further dissection of
the single specimen).

Relationships

As there remain a number of Papuasian
trigoniuline genera of which the gonopod structure
is unknown or only poorly known, I am unable to
this time to suggest any candidate sister-group for
Austrostrophus. The singular form of the anterior
gonopods is not approached in any genus known to
me, nor the presence of a modified metazonal edge.
Curiously, the posterior gonopods offer no
corresponding pecularities, and in fact these
appendages tend to be conservatively quite similar
through the subfamily.

ACKNOWLEDGEMENTS

I am very much indebted to my colleague Mark
Harvey (WAM), who on realizing that a
trigoniuline from Western Australia would be of
particular interest, brought the specimen to my
attention. I am much indebted to Dr. Harvey for the
the opportunity to examine and describe this
curious, disjunct spiroboloid, and to Patrick
Brannon, VMNH technician, for the SEM
photography.

REFERENCES

Jeekel, C. A. W. (2001). A bibliographic catalogue of the
Spirobolida of the Oriental and Australian regions
(Diplopoda). Myriapod Memoranda 4: 5-103.

Hoffman, R. L. (1994). Studies on spiroboloid millipeds.
XVIII. Speleostrophus nesiotes, the first known
troglobitic spiroboloid milliped, from Barrow Island,
Western Australia (Diplopoda: Pachybolidae:
Trigoniulinae). Myriapodologica 3: 19-24.

Schmidt, D. (1962). Uber die taxionomische Wertigkeit
von Strukturen des Metazonit-Hinterrandes bei
Diplopoden. Senckenbergiana biologia 43: 65-80.
Verhoeff, K. W. (1937). Ueber einige neue Diplopoden
aus Australien. Records of the Australian Museum 20:
133-149.

Manuscript received 11 November 2002; accepted 11 December
2002

Records of the Western Australian Museum 22: 23^5 (2003).

Psammophilous halacarids (Halacaridae, Acari) from Dampier, Western
Australia. Description of species and faunal comparison of the
mesopsammal halacarid fauna of western and eastern Australia

Ilse Bartsch

Forschungsinstitut Senckenberg, c/ o DESY, Notkestr. 31, 22607 Hamburg, Germany
e-mail bartsch@meeresforschung.de

Abstract - Twelve species of halacarid mites were extracted from tidal and
subtidal sandy deposits and coral fragments in the tropical Dampier area.
Western Australia. The genera and number of species (in parentheses) are:
Actacarus (2), Anomalohalacarus (1), Arhodeoporus (1), Copidognathus (1),
Scapthognathides (2), Scapthognathus (1)/ and Simognathus (4). All species are
diagnosed and four species, new to science, are described, viz. Actacarus
festivus sp. nov., Anomalohalacarus dampierensis sp. nov., Simognathus salebrosus
sp. nov., and Simognathus tener sp. nov. Seven of the twelve species are known
from the Indo-Pacific region, from warm-temperate Western Australia,
tropical eastern coast of Australia, Hawaiian Islands, and/or off Chile.

Key words: Australia, marine mites, psammobionts, new species, faunal
comparison

INTRODUCTION

Sandy deposits with medium and coarse grain
fractions provide a system with a large surface,
different-sized void volumes and regular interstitial
water replenishment but shelter from extreme
amplitudes of environmental parameters. These
deposits contain a specialized fauna, the interstitial
fauna or mesopsammon, terms introduced by
Nicholls (1935) and Remane (1940).

The first records of mesopsammic halacarid
species from Australia were from Rottnest Island,
Western Australia, an island with a variety of
sandy deposits. About 30 psammophilous
halacarid mites from intertidal and subtidal sandy
deposits have been described (Bartsch 1993). Otto
(1999a, b, 2000a-e, g, 2001a-c), Otto and Bartsch
(1999) and (Bartsch 2000) added more than 50 new
records to the list of Australian species extracted
from sandy deposits on the northeastern coast of
Australia, many of them being psammophilous.
Examination of both tidal and subtidal sediments
collected near Dampier, tropical Western
Australia, resulted in both new records and new
halacarid species.

STUDY AREA, MATERIAL AND METHODS

Dampier, on the Burrup Peninsula, lies in the
tropical north of Western Australia, at about 20°S,
117°E. The shoreline is characterized by rocky
platforms, rocks and mangroves. Areas with sandy

deposits with medium and coarse sand are rare.
Subtidal deposits immediately off Dampier are rich
in organic material and typical arenicolous
halacarids are generally lacking, but around the
outer islands of the Dampier Archipelago there are
coarse sediments with a psammophilous
meiofauna.

The mites were extracted from the sand by
frequently stirring with water and decanting
through a sieve with 100 pm mesh diameter. The
mites were cleared in lactic acid and mounted in
glycerin jelly. Type and voucher specimens are
deposited in the Western Australian Museum, Perth
(WAM).

Abbreviations used in the descriptions: AD =
anterior dorsal plate; AE = anterior epimeral plate;
ds-1 to ds-6 = first to sixth dorsal idiosomatic
seta(e); GA = genitoanal plate; GO = genital
opening; OC = ocular plate(s); P = palp, P-1 to P-4 =
first to fourth palpal segment; pas = parambulacral
seta(e); PD = posterior dorsal plate; PE = posterior
epimeral plate(s); pgs = perigenital setae; sgs =
subgenital setae. The legs are numbered I to IV, leg
segments 1 to 6 are trochanter, basifemur,
telofemur, genu, tibia, and tarsus.

Except one species recently described and figured
on the base of specimens from Dampier (Bartsch
2003b), the species are diagnosed and illustrated, in
general relying on individuals from the Dampier
area. In the bibliography of a given species only the
most relevant descriptive papers are cited.

24

I. Bartsch

SYSTEMATICS

Subfamily Actacarinae Viets, 1939

Genus Actacams Schulz, 1937

Actacarus festivus sp. nov.

Figure 1A-H

Material Examined

Holotype

Female, sponge garden, West of Rocky Head,
20°32.1'S, 116°26.7’E, Enderby Island, Dampier
Archipelago, Western Australia, Australia; 2-13 m,
unsorted sandy deposits, 3 August 2000, coll. C.
Bryce (WAM T52095).

Etymology

Specific name from festivus, Latin, nice, festive.
Diagnosis

Idiosomal length 205 pm. Dorsal and ventral
plates uniformly porose. Opposing margins of AD
and PD and AE and GA truncate. Length of PD less
than twice that of AD. Setae ds-2 within striated
Integument. PE with three setae; one pair of setae
within striated integument. Anterolateral margins
of GA converging. Ovipositor extending beyond
GA. Gnathosoma 0.28 of length of idiosoma.
Rostrum extending slightly beyond middle of P-2.
Tectum arched. Tibia I with three slender ventral
setae, tibiae II to IV each with one faintly
bipectinate seta. Claws on tarsi II to IV with long,
coarse tines.

Description

Female

Idiosoma. Length 205 pm, width 115 pm. Dorsal
plates uniformly and minutely porose; with few
striae of membraneous integument between
opposing margins of AD and PD. AD wider than
long (Figure 1A), length 67 pm, width 88 pm;
anterior margin arched, posterior margin almost
truncate. Gland pores in lateral margin. OC
triangular, length 15 pm, width 7 pm; with gland
pore and pore canaliculus. Length of PD 127 pm,
width 90 pm; its length 1.9 times that of AD. PD
with two pairs of gland pores, anterior pair
somewhat posterior to level with insertion of leg IV.
Dorsal setae small, pair of ds-1 slightly posterior to
the level of gland pores. Pair of ds-2 in striated
integument; ds-3 to ds-5 on PD, arranged as
illustrated. Pair of ds-5 level with gland pores. Pair
of ds-6 on anal plate.

Length of AE 91 pm, width 103 pm; with three
pairs of setae; posterior margin of AE truncate
(Figure IB). PE extending beyond insertion of leg
IV. One pair of setae within striated integument

between PE. Length of GA 87 pm, width 72 pm;
anterior margin truncate, lateral margin diverging
from anterior margin to level with posterior edge of
PE. GA with three pairs of pgs. Ovipositor long,
extending beyond anterior margin of GA and
posterior margin of AE.

Gnathosoma. Length 59 pm, width 46 pm, 1.2 times
longer than wide. Gnathosoma 0.28 of length of
idiosoma. Tectum broadly arched (Figure ID).
Rostrum short, far from reaching end of P-2. P-2
with a dorsal seta. P-3 with spiniform medial
process. P-4 with three setae in basal whorl, one
spur, one ventral and lateral seta, and a spur-like
tip (Figure 1C).

Legs. Leg IV longer than leg I. Lateral flanks of
legs I and II delicately punctate. Tibia I with slender
base, then widened; its length surpassing that of
telofemur I. Tibiae and relevant telofemora of
following legs similar in length. Telofemur I 1.8
times longer than high. Leg chaetotaxy (solenidia
included, pas excluded): leg I, 0, 2, 3, 5, 7, 7 (Figure
IE); leg II, 0, 2, 3, 4, 5, 4 (Figure IF); legs III and IV,
1, 2, 2, 3, 5, 3 (Figures 1G and H). Genu II with
dorsal alveolus but no seta seen. Tibia I with three
slender ventral setae, a ventrolateral seta and three
dorsal setae. On tibia II ventromedial seta slightly
wider than ventral seta. Ventromedial seta on tibia
III long and wide, apical seta of tibia III only slightly
widened. Ventromedial seta of tibia IV delicately
widened, apical seta slender. Tarsi I to IV each with
pair of pas.

Length of claws increasing from tarsus I to III.
Claws on tarsus I with accessory process but no
pecten. Pectines of claws II to IV with very coarse
tines. Central sclerite with small bidentate process.

Remarks

Actacams festivus has a short PD relative to the
length of the AD, there is one pair of setae within
the membraneous ventral integument, and the
margins of the female GA are convergent instead of
almost parallel-sided. The characters are shared
with A. giganteus Krantz, 1971, a species recorded
from the Caribbean (Krantz, 1971). The Caribbean
species is somewhat larger, its idiosoma and the
dorsal and ventral plates are more slender than in
A. festivus, the dorsal plates are coarsely punctate in
A. giganteus but minutely porose in A. festivus.

The arrangement of the dorsal plates resembles
that of A. latus Newell, 1984. Beside the number of
setae on the AE (three in A. festivus, four in A. latus)
there are differences in the size (length of female
205 pm in A. festivus, 334-355 in A. latus), the
ornamentation of the dorsal plates (minutely and
uniformly porose in A. festivus, pitted in A. latus),
shape of the female GA and the arrangement of the
pgs. Actacarus latus is recorded from the Eastern
Pacific, from off the coast of Chile (Newell, 1984).

The shape of the female GA of A. australis Bartsch,

Psammophilous halacarids from Western Australia

25

1993 is similar to that of A.festivus. In contrast to A.
festivus the PE of A. australis bears three ventral
setae and the ovipositor is much shorter.

Distribution

Indian Ocean, Western Australia, Dampier. From
shallow water deposits.

Actacarus pacificus Bartsch, 1979
Figure 2A-D

Actacarus pacificus Bartsch, 1979: 231-234, figures 1-
14; Bartsch, 1993: 75, 76, figure 1; Abe, 1997: 33-
34, figure 2A-C.

Actacarus orthotectus Newell, 1984: 245-247, figures
705-707.

Actacarus marindicus Otto, 2000c: 116-119, figures
3a-e, 4a-e (new synonymy).

Material Examined

Australia: Western Australia: 1 female, Dampier,
20°39'S, 116°42'E, sandy beach, coarse, unsorted
sand in middle part of tidal slope, 0-3 cm sediment
depth, 3 August 2000, coll. I. Bartsch (WAM
T52096).

Diagnosis

Idiosomal length 263 pm. Dorsal and ventral
plates with delicate punctation. Opposing margins
°f AD and PD and AE and GA truncate (Figure
2A). PD more than twice the length of AD. OC with
triangular anterior portion extending beyond gland
pore. Pair of ds-2 within membraneous integument.

26

I. Bartsch

Figure 2 A-D. Actacarus pacificus Bartsch, 1979, female. A, Idiosoma, dorsal; B, idiosoma, ventral; C, gnathosoma,
ventral; D, leg I, medial. Scale = 50 pm.

AE with four pairs of setae. PE with one dorsal and
two ventral setae (Figure 2B). Ovipositor in rest
extending beyond anterior margin of GA.
Gnathosoma 0.27 of idiosomal length. Rostrum
about as long as gnathosomal base. Tectum very
slightly arched. Rostrum extending almost to end of
P-2 (Figure 2C). Both pairs of maxillary setae in
distal half of rostrum. Legs less than 0.7 of
idiosomal length. Genua shorter than telofemora
and tibiae. Leg chaetotaxy: leg I, 0, 2, 3, 5?, 7, 7; leg
II, 0, 2, 3, 4, 5, 4; legs III and IV, 1, 2, 2, 3, 5, 3. Tibia I
with three slender ventral setae (Figure 2D); tibiae
II to IV with two ventral setae, on tibia III these
setae faintly bipectinate. Paired claws on tarsus I
with accessory process but no pecten, claws II to IV
with few large tines.

Remarks

According to Bartsch (1993) and Abe (1997),
Actacarus pacificus is found widely spread in the
Indo-Pacific region. In contrast, Otto (2000c)
expected specimens from Rottnest Island, Western
Australia, collected and identified by Bartsch (1993),
not to be identical with A. pacificus. Otto described
the Australian specimens under the name of
Actacarus marindicus Otto, 2000. The differences
between the individuals from the Hawaiian Islands
and Australia were said to be (1) a coarser
punctation of the PD in the Hawaiian material and
(2) presence of a single bipectinate seta on tibia III
of the type specimens, instead of two such setae as
present in the Australian individuals. In respect to
the first mentioned character, the differences stated
by Otto (2000c) cannot be confirmed by the present

author. The dorsal plates of the holo- and paratype
female and male of A. pacificus are delicately
punctate, as the ventral plates are. Unfortunately,
Otto (2000c) did not mention if the illustration
(Otto, 2000c: figure 8b) was prepared on the base of
the holo- or a paratype, a female or male. In the
illustration the position of the setae assumedly
representing the ds-4 is unusual (and not
corresponding to the situation in either the holotype
or the paratypes), and the ds-3 are not illustrated.
The second character used by Otto (2000c) to
distinguish between A. pacificus and A. marindicus
was the shape of a seta on tibia III. The basal one of
the two ventral setae on tibia III is longer than the
distal one and delicately bipectinate. The distal seta
has fuzzy margins and such a tip (Otto, 2000c: 118)
due to a very subtle bipectination. These fuzzy
margins are not recognizable when viewing the
edge of that seta. Presence or absence of fuzzy
margins is inadequate for discriminating between
species as their perceptibility is influenced by the
preservation, storing, clearing and mounting
medium.

Bartsch (1993: 76) mentioned the shape of the
basal seta on tibia I as a possible diagnostic
character to distinguish between the Hawaiian and
Western Australian specimens. A re-examination of
the holo- and paratypes showed that the relevant
seta is somewhat more stout than the other ventral
setae but not distinctly pectinate.

Otto (2000c) also discussed the state of A.
orthotectus Newell, 1984. That species was described
from the Juan Fernandez Islands, Chile (Newell,
1984). Bartsch (1993) and Abe (1997) believed it to

Psammophilous halacarids from Western Australia

27

be synonymous with A. pacificus. Otto (2000c) re-
established the Chilean species (A. orthotectus)
because of its slender P-2 and presented an
illustration of the palps of both a Chilean and
Western Australian specimen (Otto, 2000c: figure 3e
and f). The palps are said to show a lateral aspect.
In general, the P-2 of Actacarus species are slightly
flattened but narrow in dorsal aspect. It is likely
that one of the illustrations (Otto, 2000c: figure 3f)
represents a dorsolateral aspect of the palps, a
notion corroborated by the insertion of the dorsal
seta in that figure. In lateral aspect the shape of the
palps may be the same in specimens from both the
southeastern and northern Pacific.

Nonetheless, several cryptic species may exist in
the Indo-Pacific region, each restricted to a small-
scale geographical area, and the specimens from
Rottnest Island (A. marindicus) may prove to be
distinct. The presently described morphological
characters allow no separation between individuals
from the Hawaiian Islands and Western Australia.

The Actacarus species recorded from Australia are
A. australis, A. chelonis Otto, 2000, A. cornutus Otto,
2000, A. nanus Otto, 2000, A. pacificus, A. spinosus
Otto, 2000, and the above described A. festivus. The
AE of each of A. pacificus, A. nanus and A. spinosus
bears four pairs of setae, the PE two ventral setae.
A. spinosus, the smallest species, has an idiosomal
length of 186-194 pm, its P-4 has a conspicuously
bent seta (instead of a spur), on tibia I is one of the
ventral setae coarsely bipectinate, one faintly
pectinate, tibiae III and IV both bear one long
bipectinate and one very short and slender ventral
seta. Actacarus nanus has an idiosomal length of
198-212 pm, the ds-2 are close to the level of the
posterior margin of the AD and the OC are rather
short, the female GA is short, the distance from the
anterior margin of the GO to the GA almost equals
half the length of the AE, and none of the ventral
setae on tibia I is bipectinate. Actacarus pacificus has
an idiosomal length of 235-282 pm (Australian
specimens), the length of the GA relative to that of
the AE is less than in A. nanus, the ventral setae on
tibia I are apparently smooth.

Distribution

Southeastern Pacific (Chile, Robinson Crusoe
Island), Northeastern Pacific (Hawaiian Islands),
Sea of Japan (Hokkaido), Eastern Indian Ocean
(Western Australia, Rottnest Island, Dampier).
Intertidal to 10 m.

Subfamily Anomalohalacarinae Bartsch, 1985

Genus Anomalohalacarus Newell, 1949

Anomalohalacarus dampierensis sp. nov.
Figures 3A-H, 4A-G

Material Examined

Holotype

Male, Dampier, 20°39'S, 116°42'E, Western
Australia, Australia; sandy beach, coarse, unsorted
sand in middle part of tidal slope, 0-3 cm sediment
depth, 3 August 2000, coll. I. Bartsch (WAM
T52097).

Paratype

Australia: Western Australia: 1 protonymph,
same data as for holotype (WAM T52098).

Etymology

An adjective referring to the type locality
Dampier.

Diagnosis

Idiosomal length 300 pm. AD and PD almost
equal in length. PD undivided. Setae ds-5 lacking.
Setae on epimera IV lacking. Anterior margin of
male GA arched. GA with at least four pairs of pgs.
P-2 with two setae. Tibia I with two short,
bipectinate ventral spurs. Tibia I with eight setae,
two of them spur-like. Tibiae II, III and IV each
with pair of ventral setae which are slightly
pectinate on tibiae III and IV but pectinate and
slender on tibia II. One of the two setae on
telofemur IV conspicuously stout.

Description

Male

Idiosoma. Length 300 pm; very slender, much
longer than wide. Dorsal and ventral plates delicate.
Membraneous integument faintly striated. Length
of AD 57 pm, width 39 pm; with pair of gland pores
in lateral margin and pair of ds-1 posterior to gland
pores (Figure 3A). PD undivided, its length 57 pm,
width 30 pm; with pair of pores in posterior third of
plate and small ds-6 in posterior margin. AD and
PD each with pair of gland pores. Setae ds-1 to ds-4
long, slender; ds-1 inserted on AD, ds-2 and ds-3
within striated integument; ds-5 not seen; ds-6 very
short, in posterior margin of PD.

AE divided by striated integument into right and
left half (Figure 3B). Length of AE 42 pm (to
camerostome). Each half with three setae and an
epimeral pore. Long apodemes between epimera I
and II. PE in posterior fourth of idiosoma. Each
plate with one dorsal and two ventral setae; no seta
in area representing epimeral plate IV. Anterior
margin of GA broadly arched, with at least four
pairs of perigenital setae (genital region obscured
by legs III and IV and hence setae difficult to
discern). Spermatopositor extending far beyond
GA. Position of sgs not discemable in the single
male available.

Gnathosoma. Very slender (Figure 3C), its length

28

I. Bartsch

87 jam, width 28 jam. Rostrum almost as long as
gnathosomal base; with two pairs of long maxillary
setae; apex with two pairs of minute rostral setae.
P-2 with short, slender basal and stout distal seta
(Figure 3D). P-3 with large medial spine. P-4 with
three setae in basal whorl; apex with minute ventral
and lateral seta and flattened dorsal seta.

Legs. Leg I longer and wider than the other legs.
Genu and tibia I almost equal in length and shorter
than telofemur I. Both lateral and medial fossa
membrane of tarsus I large. Tarsi II to IV lack
prominent membranes. Leg chaetotaxy (solenidia
and pas included, famulus excluded): leg I, 1, 2, 3,
5, 8, 9; leg II, 1, 2-3, 3, 5, 5, 6; leg III, 1, 1, 2, 3, 5, 5;
leg IV, 0, 0, 2, 3, 5, 5. Dorsal seta of basifemur I
conspicuously long and stout (Figure 3G). Genu I
with two short, slightly bipectinate spurs, inserted
adjacent and ventromedially. Tibia I with two

ventral spurs; most of dorsal setae slender except
rather thick distolateral seta. Tarsus I with three
ventral setae, one spur-like, two short and slender.
Dorsal and dorsomedial fossary seta slender,
dorsolateral fossary seta and dorsolateral solenidion
conspicuously thick (Figure 3E). Famulus slender,
pressed to lateral fossa membrane; its length 8 pm.
Genua II to IV with slender ventral setae. Tibiae II
to IV each with pair of ventral setae; on tibia II one
of these setae pectinate and one slender (Figure 3H),
on tibiae III and IV both setae equal-sized, with very
delicate pectination. Tarsus II with large, claviform
solenidion in dorsolateral position (Figure 3F); the
three dorsal setae slender. Tarsi I to IV with pair of
pas singlets. Telofemora III and IV each with two
dorsal setae, on telofemur IV one of these setae
stout.

Paired claws of tarsus I short, with basiventral

Psammophilous halacarids from Western Australia

29

Figure 4 A-G, Anomalohalacarus dampierensis sp. nov., protonymph. A, Idiosoma, dorsal; B, idiosoma, ventral; C, leg I,
medial; D, tip of tarsus I with claw; E, leg II, medial; F, leg III, ventral; G, trochanter, femur, genu IV. Scale =
50 pm (A-C, E-G), scale = 10 pm (D).

process (as in protonymph. Figure 4D). Paired
claws of tarsi II to IV with accessory process but
without pecten. Central sclerite with minute claw-
like process.

Protonymph

Idiosomal length 268 pm, width 100 pm.
Arrangement of plates and setae as illustrated
(Figure 4A and B). AE as in male with three pairs of
setae, PE with one dorsal and two ventral setae.
Gnathosoma slender, length 80 pm, width 25 pm,
number and arrangement of setae same as in adults.
Leg chaetotaxy (with solenidia and pas): leg I, 1, 2,
3, 5, 7, 9 (Figure 4C); leg II, 1, 2, 3, 4, 5, 6 (Figure 4E);
leg III, 1, 1, 2, 3, 5, 5 (Figure 4F); leg IV, 0, 0+2 (basi-
+ telofemur), 3, 5, 5. Genu I with two spur-like,
pectinate setae. Tibia I with two spur-like setae.
Tibiae II with one slender and one pectinate seta;
tibiae III and IV each with two faintly bipectinate
setae. One of dorsal setae on telofemur IV stout
(Figure 4G).

Remarks

Most species of the genus Anomalolmlacarus are
known from the North Atlantic and the
Mediterranean. There is a single species from the
western Pacific, from Japan, and three from the
Indian Ocean, two from Western Australia and one
species of unknown identity from the Bay of Bengal
(Abe, 1996; Bartsch, 1976a, b, 1981b, 1985a, 1991a,
1993; Monniot, 1967; Morselli and Mari, 1981, 1982,
1989; Rao, 1970).

The idiosoma and dorsal and ventral plates are
similarly shaped in the males of the two Australian
species, A. macellus Bartsch, 1993, from Rottnest
Island, and A. dampierensis. The two species differ
in the setation of the legs. Tibia I of A. dampierensis
bears eight setae, two of them are spur-like, in
contrast, tibia I of A. macellus has 10 setae. The
protonymph of A. dampierensis has a longer genital
plate than that nymph of A. macellus.

Anomalohalacarus biformis Abe, 1996, from
Hokkaido, Japan (Abe, 1996), has 10 setae on tibia I.

30

I. Bartsch

The species is easily separated from the two
Australian ones by the three ventral bristle-like
setae on tibiae II and III.

Fourteen species have been described from the
North Atlantic and adjacent basins. Two species,
Anomalohalacarus anomalus (Trouessart, 1894) and A.
litoralis Bartsch, 1981, differ from the others in that
the idiosomal integument is very prominently
striated (cf. Monniot, 1967: figure 1C), the P-2 bears
two large setae, one in the basal, one in the distal
portion, and tibia I three ventral setae (two spur-
like, one seta-like) in the basal half of the segment.
In the other North Atlantic species, the striae of the
integument are rather delicate, the P-2 bears a single
seta which is slender and inserted near the base of
that segment, and in the basal portion of tibia I
there are one or two setae which are either setiform
or seti- and spiniform. Except for A. marcandrei
Monniot, 1967, the PD of adult North Atlantic
species is divided into a right and left half.

The three species from the Indo-Pacific region
differ from those of the North Atlantic and its
adjacent basins by (1) absence of the ds-5 and (2)
absence of the posterior pair of setae on the PE
(generally attributed to epimeron IV). The PD is
undivided, the idiosomal integument delicately

striated, P-2 bears two setae, the basal seta is short
and slender, similar to that of the majority of the
North Atlantic species, the posterior seta is stout,
resembling that seta of A. anomalus and A. litoralis.
In the basal portion of tibia I there are more than
two ventral setae.

Distribution

Indian Ocean (Western Australia, Dampier).
Intertidal.

Subfamily Halacarinae Viets, 1927

Genus Arhodeoporus Newell, 1947

Arhodeopoms corallicolus Otto, 2000
Figures 5A-F, 6A-F

Arhodeoporus corallicolus Otto, 2000g: 5-7, figure 4.

Material Examined

Australia: Western Australia: 1 female, 1 male,
Dampier Archipelago, Enderby Island, West of
Rocky Head, 20°32.1'S, 116°26.7'E, sponge garden,
2-13 m, unsorted sandy deposits, 3 August 2000,
coll. C. Bryce (WAM T52099 and T52100).

Figure 5 A-E. Arhodeoporus corallicolus Otto, 2000. A, Idiosoma, dorsal, male; B, idiosoma, ventral, male; C, genitoanal
plate, male; D, gnathosoma, ventral, male; E, genitoanal plate, female; F, palp, lateral, male. Scale = 50 pm (T,
tectum).

Psammophilous halacarids from Western Australia
Diagnosis

Idiosomal length 400M11 pm. Surface of plates
smooth but integument pierced by numerous
canaliculi. Outline of such porose areolae as
illustrated (Figure 5A and B). OC with long tail
and one small cornea and, beneath, small spots of
eye pigment. Anterior margin of PD ovate. Each of
the four pairs of gland pores with a small tube.
Pair of ds-2 within striated integument; ds-3 on
small platelets; ds-5 just posterior to gland pore.
Punctation of AE coarser than that of dorsal plates
and GA. Opposing margins of AE and GA
truncate. AE with pair of epimeral pores. PE with
one dorsal and three ventral setae. Female GA
with three pairs of pgs; genital sclerites with three
pairs of sgs; two pairs of internal genital acetabula.
Ovipositor extending slightly beyond the level of
anterior pair of pgs (Figure 5E). Male GA with 17

31

pgs; genital sclerites with five pairs of sgs and a
slightly projecting posterior cone (Figure 5C).
Spermatopositor extending well beyond
anteriormost pair of pgs. Gnathosoma slender,
almost three times longer than wide and 0.44 of
idiosomal length. Rostrum distinctly longer than
gnathosomal base (Figure 5D). The latter with
marginal porose areolae. Rostral trough long;
truncate tectum in basal portion of gnathosomal
base (Figure 5D). Palps slender (Figure 5F). Basi-
and telofemora II to IV flattened; telofemora II to
IV about twice as long as high; basifemora dorsally
slightly projecting. Tibiae slender, cylindrical.
Fleight of telofemora II to IV 1. 8-2.0 times that of
relevant tibiae. Tibia I longer than telofemur I,
tibiae of following legs as long as or shorter than
telofemora. Leg chaetotaxy (solenidia included,
pas excluded): leg I, 1, 2, 3, 4, 8, 7 (Figure 6A); leg

Figure 6 A-F. Arhodeoporus corallicohis Otto, 2000, male. A, Leg I, medial; B, leg II, medial; C, leg III, medial; D,
basifemur to tarsus IV, medial; E, leg III, medial; F, leg IV, medial. A-D, male from Dampier; E and F,
paratype from Great Barrier Reef. Scale = 50 pm.

32

I. Bartsch

II, 1, 2, 3, 4, 7, 4 (Figure 6B); leg III, 1, 2, 2, 3, 5, 4
(Figure 6C); leg IV, 0, 2, 2, 3, 5, 3 (Figure 6D).
Ventral flank of tibia I with one short, pectinate
and three long, smooth setae. Tibiae II to IV with
2, 1, 1 bipectinate and 1, 1, 1 smooth and slender
seta. Tarsus I short, less than half length of tibia I.
Tarsi I and II with slender pas; tarsus III with
slender medial and short, wide lateral pas; tarsus
IV with pair of short and wide pas. Claws with
accessory process and pecten; tines of pectines
very short (seen only at high magnification and oil
immersion).

Remarks

The specimens from the Dampier Archipelago
have conspicuously flattened telofemora II to IV,
their height is almost twice that of the tibiae. The
height of the telofemora of the Great Barrier Reef
specimens (Otto 2000g: figure 4f and present paper:
Figure 6E and F) is less striking, 1.5-1. 7. The fixative
and the contraction of the muscles may influence
the shape of the telofemora. The slight difference in
the shape of the PD is expected to be within the
variability of this character; the anterior portion of
the holotype PD is broadly rounded (Otto 2000g:
figure 4a), that of the specimen from Dampier more
slender (Figure 5A).

Species most similar to Arhodeoporus
corallicolus are A. caudatus Otto, 2000, A.
clypeatus Otto, 2000, A. lizardensis Otto, 2000, and
A. psammophilus Bartsch, 1993, all from
Australia, from the Great Barrier Reef, the
Queensland Plateau and from Rottnest Island,
off Perth (Otto, 2000g; Bartsch, 1993). Further
species are A. mactanus Bartsch, 1991, from the
Philippine Island Mactan, and A. longirostris
Bartsch, 1981, from off the Isles Glorieuses,
Mozambique Channel (Bartsch, 1981a, 1982,
1991b). Arhodeoporus corallicolus can be identified
on the base of the combination of: OC with one
cornea; AE and GA separated; punctation of AE
coarser than that of the other plates; tibia I with
short, delicately pectinate seta; basi- and
telofemora flattened. An unique character is that
the female genital sclerites bear three pairs of
sgs, in contrast to two pairs of setae which is the
most common state of this character.

Arhodeoporus corallicolus has very flattened
telofemora. In general, the telofemora of halacarid
mites are almost cylindrical, but in many species
the legs bear large ventral lamellae, e.g. in species
of the Copidognathus gibbus group (Bartsch, 1994b;
Otto, 2000f). In A. corallicolus there are no such
lamellae.

Distribution

Great Barrier Reef, northeastern Australia, and
Dampier, northwestern Australia. Inhabitant of
shallow water sandy deposits.

Subfamily Copidognathinae Bartsch, 1983
Genus Copidognathus Trouessart, 1888
Copidognathus meridianus Bartsch, 2003

Copidognathus meridianus Bartsch, 2003b: xx-xx,
figures 1A-H, 2A-H, 3A-F.

Material Examined

Australia: Western Australia: 3 females, 1 male,
Dampier, 20°39'S, 116°42'E, unsorted sand in mid-
and upper tidal shore, 31 July 2000 and 1 August
2000, coll. I. Bartsch (WAM T45195-T45198); 1
female, north of Cape Preston, 40 Mile Beach, coarse
unsorted sediment, upper tidal shore, 6 August
2000, coll. I. Bartsch (WAM T45199).

Diagnosis

Idiosomal length 360-385 pm. Dorsal plates
almost uniformly ornamented with small pits
arranged within polygons. AD with three delicately
raised areolae. PD without prominent costae. OC
with minute cornea. PD with pairs of pores at the
level and posterior to ds-5; each pore with large
internal gland. Female GA with four pairs of pgs;
male GA with 15-16 widely scattered pairs of pgs.
Length of gnathosoma almost one third of that of
idiosoma. Length:width ratio of gnathosoma 1:0.7;
rostrum triangular, about as long as gnathosomal
base and extending just beyond P-2. Tectum
truncate. Leg chaetotaxy (solenidia and pas
included): leg I, 1, 2, 5, 4, 7, 11; leg II, 1, 2, 5, 4, 7, 8;
leg III, 1, 2, 2, 3, 5, 5; leg IV, 0, 2, 2, 4, 5, 5. Tibiae II
to IV with 2, 1, 1 bipectinate setae. Tarsi I to IV with
4, 4, 3, 3 dorsal setae (solenidia included). Pectines
of claws II to IV with delicate tines.

Remarks

This rather large-sized psammophilous
Copidognathus species can be separated from
congeners by the unique combination of (1) dorsal
plates almost uniformly ornamented and (2) genu
IV with four setae.

Distribution

Tropical Western Australia, Dampier. Present in
sandy deposits, in mixed coarse and fine sand. In
the Dampier area, this species dominated the
intertidal psammophilous halacarid fauna.

Subfamily Lohmannellinae Viets, 1927

Genus Scaptognathides Monniot, 1972

Scaptognathides haivaiiensis Bartsch, 1988
Figure 7A-E

Scaptognathides haivaiiensis Bartsch, 1988: 221, 222,
figures 27-30; Bartsch, 1991c: 59, figure 1A-H.

Psammophilous halacarids from Western Australia

33

Figure 7 A-D. Scaptognathid.es hawaiiensis Bartsch, 1988, female. A, Idiosoma, dorsal; B, idiosoma, ventral; C,
gnathosoma, ventral; D, ovipositor; E, leg I, medial, (ept, epimeral tube). F-K. Scaptognathides ornatus Bartsch,
1988, female. F, gnathosoma, ventral; G, leg I, ventral; H, tip of tarsus II, medial; I, genital opening; J,
idiosoma, dorsal; K, idiosoma, ventral. Scale = 50 pm (A-G, I-K), scale = 10 pm (H).

Material Examined

Australia: Western Australia: 3 females,
Dampier, sandy beach, coarse, unsorted sand, lower
part of tidal slope, 2-15 cm sediment depth, 4
August 2000, coll. I. Bartsch (WAM T52101 and
T52102).

Diagnosis

Idiosomal length 170-180 pm. Dorsal and ventral
plates minutely pitted. AD, OC and PD each with
one pair of gland pores (Figure 7A). Pairs of ds-1,
ds-2 and ds-3 on AD, ds-4 and ds-5 on PD, adanal
setae on anal valves. AE with three pairs of setae

34

I. Bartsch

and pair of small epimeral tubes (Figure 7B). GA
with two pairs of pgs, genital sclerites with one pair
of sgs. Gnathosomal length 0.44-0.47 times of that
of idiosoma. Margin of tectum truncate. Rostrum
slightly longer than gnathosomal base (Figure 7C).
Two of the four apical palpal spines pectinate. Leg I
somewhat longer and considerably more stout than
following legs. Length of legs I and IV about 0.75 of
that of idiosoma. Leg chaetotaxy (very short
solenidia excluded, pas included): legs I and II, 1, 2,

4, 4, 5, 5; legs III and IV, 1, 2, 2, 3, 5, 5. Length:height
ratio of telofemora II to IV about 2. 2-2.3. Paired
claws of tarsus I with large, umbrella-like arranged
tines (Figure 7E). Pectines of claws of tarsi II to IV
seen only at higher magnification; basalmost tine
twice the length of the other very delicate tines.

Supplementary notes

Ovipositor with 10 pairs of delicately sclerotized,
lamellar genital spines (Figure 7D).

Remarks

The PD of Scaptognathid.es hawaiiensis bears a
single pair of gland pores situated near the
posterior margin of the plate. In the other
Australian species, the PD has two pairs of gland
pores, one pair about the level of insertion of leg IV,
the other pair near the posterior margin, similar to

5. hawaiiensis.

Records of S. hawaiiensis are from the Hawaiian
Archipelago, Hong Kong and northwestern
Australia. There are small differences in the shape
of the female GA, with the anterior margin oviform
in the individuals from the Hawaiian Archipelago
(Bartsch 1988: figure 28) but almost truncate in
those from the Dampier area. Considering the
delicateness of the plates, which easily can be
deformed, the differences mentioned are thought to
be negligible.

Distribution

Northeastern Pacific (Hawaiian Islands), South
China Sea (Hong Kong), Eastern Indian Ocean
(Western Australia, Dampier). Intertidal sandy
deposits.

Scaptogtiathides omatus Bartsch, 1988
Figure 7F-K

Scaptognathides omatus Bartsch, 1988: 222-224,
figures 31-39.

Material Examined

Australia: Western Australia: 1 female, Dampier,
20°39'S, 116°42'E, sandy beach, coarse, unsorted
sand in lower part of tidal slope, 2-15 cm sediment
depth, 4 August 2000, coll. I. Bartsch (WAM
T52103).

Diagnosis

Idiosomal length 217 pm. Dorsal plates
reticulated (Figure 7J). AD with one pair of pores,
OC and PD each with two pairs of gland pores.
Pair of ds-1, ds-2 and ds-3 on AD, ds-4 near
anterolateral comers of PD, ds-5 situated in middle
of plate anterior to the level of pair of glp-4.
Marginal areas of ventral plates faintly reticulated,
ventral portions punctate. GA with two pairs of
pgs (Figure 7K) and two pairs of internal genital
acetabula (Figure 71), its genital sclerites with one
pair of sgs. Length of gnathosoma 0.48 times that
of idiosoma. Tectum concave, median margin
truncate. Rostrum somewhat longer than
gnathosomal base (Figure 7F). Two of apical palpal
spines pectinate. Legs I and IV almost 0.75 of
idiosomal length. Length of genu I about 0.6 and
0.2 times that of tibia and telofemur I, respectively.
Telofemora II to IV slender, almost three times
longer than high. Leg chaetotaxy (very short
solenidia excluded): legs I and II, 1, 2, 5, 4, 5, 5;
legs III and IV, 1, 2, 2, 3, 5, 5. Paired claws of
tarsus I with long, umbrella-like arranged tines
(Figure 7G); claws of tarsi II to IV with minute
accessory process, pecten with tines, two
basalmost tines enlarged (Figure 7H).

Remarks

The two species, Scaptognathides ornatus and S.
hawaiiensis, present in Dampier in the same beach,
are easily separated even at low magnification; S.
omatus is larger, has reticulate dorsal plates and
slender telofemora II to IV.

From Australian shores, six species of
Scaptognathides are recorded, S. australis Bartsch,
1993, S. haioaiiensis, S. heraldensis Otto, 2000, S.
ornatus, S. tomkinsae Otto, 2000, and S. undulatus
Otto, 2000. Characters to identify adults (and
assumedly nymphs as well are):

Scaptognathides australis: OC slender, at least four
times longer than wide, and with a single gland
pore; PD with two pairs of gland pores.
Scaptognathides hawaiiensis: the only species with a
single pair of gland pores on OC and PD.
Scaptognathides heraldensis: OC with one gland pore,
PD with two pairs of pores; in contrast to S.
australis, the OC are rhomboid, about twice as
long as wide.

Scaptognathides omatus: both OC and PD with two
pairs of gland pores. Telofemora II to IV
slender; tarsus III with three dorsal setae.
Scaptognathides tomkinsae: PD with two pairs of
gland pores; OC with a single pore, its posterior
edge with a minute seta instead of a gland pore.
Scaptognathides undulatus: both OC and PD with two
pairs of gland pores; tarsus III with four dorsal
setae, instead of three as present in the other
species.

Psammophilous halacarids from Western Australia

35

Distribution

Northeastern Pacific (Hawaiian Islands), Eastern
Indian Ocean (Western Australia, Dampier).
Intertidal.

Genus Scaptognathus Trouessart, 1889

Scaptognathus exquisitus Otto, 2000
Figure 8A-E

Scaptognathus exquisitus Otto, 2000b: 539-543, figure
4. '

Material Examined

Australia: Western Australia: 1 female, Enderby
Island, West of Rocky Head, 20°32.1'S, 116°26.7'E,
sponge garden, 2-13 m, unsorted sandy deposits, 3
August 2000, coll. C. Bryce (WAM T52104).

Diagnosis

Idiosomal length 210 pm. Dorsal and ventral
plates pitted, AD with the pits arranged within
polygons (Figure 8A). AD very wide, posterior
margin arched. OC small, rounded. Anterior
margin of PD truncate. AE wider than long, with
large epimeral pores and circular markings. PE with
funnel-like structures anterior to insertion of legs III
and IV (Figure 8A and D). Female GA bipartite;

posterior portion with two pairs of pgs (Figure 8B);
genital sclerites with single pair of sgs. Gnathosoma
almost twice as long as wide; its length 0.79 of that
of idiosoma. Spatula-shaped rostrum shorter than
gnathosomal base (Figure 8C). Legs slender; leg I

0. 7 of idiosomal length. Lateral flanks of basi- and
telofemora with small, deep foveae. Leg chaetotaxy
(solenidia, famuli and pas excluded; bipectinate
setae in roman numerals): leg I, 1, 1, 4+II, 3+II, 4+V,
4+1 (Figure 8E); leg II, 1, 1, 5, 3+1, 3+II, 3+1; leg III, 1,

1, 2, 3, 3+III, 3; leg IV, 1, 1, 2, 3, 3+III, 3. Tarsus I
with 'hollow' claviform solenidion and solid,
digitiform famulus. On either side of tarsi I and II
one setiform and one distinctly shorter pas. Tarsus
III with pair of setiform pas, medial pas with
adjacent short seta. Tarsus IV with pair of small,
setiform pas. Claws with small accessory process,
else smooth. Central sclerite without dent-like
process.

Supplementary notes

PE with funnels anterior to insertion of legs III
and IV (Figure 8D).

Remarks

The individual in Otto (2000b: figure 4A, B) shows
much wider areas between the dorsal and ventral
plates than are present in the specimen from the

Figure 8 A-E. Scaptognathus exquisitus Otto, 2000, female. A, Idiosoma, dorsal; B, idiosoma, ventral; C, gnathosoma,
ventral; D, epimeral plate III anterior to insertion of leg III, with funnel-shaped structure; E, leg I, ventral.
Scale = 50 pm (A-C, E), scale = 10 pm (D).

36

I. Bartsch

Dampier Archipelago. Such a difference is common
within ovigerous halacarid females, depending on
the number and size of eggs. The difference in the
length:width ratio of the gnathosoma (0.66 in the
specimen from the Great Barrier Reef vs 0.79 in the
one from Dampier) most likely is due to
compression of the holotype.

Marginal cavities on the PE, anterior to the
insertions of legs III and IV, have not been
described before. They are also present in two
paratype females (Zoological Museum in Hamburg
A57/00) from the Great Barrier Reef.

Distribution

Scaptognathus exquisitus seems to be widespread
along the coast of tropical Australia; records are
from eastern Australia, from 19 to 14°S, and
western Australia, at 20°S, from sandy deposits at a
depth range of 0.5 to 10 m.

Subfamily Simognathinae Viets, 1927
Genus Simognathus Trouessart, 1889

Simognathus platyaspis Otto, 2000
Figure 9A-E

Simognathus platyaspis Otto, 2000a: 519-521, figures
14A-D, 15A-D.

Material Examined

Australia: Western Australia: 1 female, Burrup
Peninsula, Watering Cove, 20°35'S, 116°58'E, coral
block, low water edge, 1 August, 2000, coll. I.
Bartsch (WAM T52105).

Diagnosis

Idiosomal length 315 pm, width 170 pm. Dorsal
plates uniformly foveate, integument delicately
punctate. Posterior two-third of AD with brown
pigmentation. AD longer than PD (Figure 9A). OC
triangular. AD with hyaline lens, OC with cornea.
Ventral plates almost uniformly foveate except for

(1) a portion immediately posterior to camerostome,

(2) anterior portion of GA and (3) around GO
(Figure 9B). GA with four pairs of pgs. Adanal setae
in ventral position. Ventral flank of gnathosoma
foveate. Triangular lamella forming a tectum
(Figure 9C). Second palpal segment with ventral
protuberance and one long seta (Figure 9D). Tibiae
club-shaped. Leg chaetotaxy (solenidia omitted, pas
included): leg I, 1, 2, 2, 4, 5, 6; leg II, 1, 2, 3, 4, 5, 6;
legs III and IV, 1, 1, 2, 3, 5, 5. Ventral seta of tibia I
basally wide, with apophysis, then tapering (Figure

Figure 9 A-F. Simognathus platyaspis Otto, 2000, female. A, Idiosoma, dorsal (stippled line along margin of brown
pigmentation); B, idiosoma, ventral; C, part of gnathosoma, dorsal; D, part of gnathosoma, lateral; D, leg I,
medial. Scale = 50 pm.

Psammophilous halacarids from Western Australia

37

9E). Pair of ventral spines of tibiae II coarsely
pectinate, that pair of spines on tibiae III and IV
only slightly pectinate. Tarsi I and II each with three
dorsal setae, one ventral seta and pair of single
eupathid pas. Tarsi III and IV each with three dorsal
setae, one ventral seta and medial pas. Tarsus I with
large median claw and slender paired claws. Paired
claws of tarsi II to IV with accessory process, else
smooth; central sclerite with small process.

Remarks

Similarly shaped and pigmented plates and the
same outline of palps and tibiae are present in S.
maculatus Bartsch, 1994, S. specialis Otto, 2000, S.
versicolor Otto, 2000, and S. xandarus Otto, 2000, all
from Australia, in S. glareus Bartsch, 1985 from New
Zealand, in S. latitarsus Proches, 2002 from South
Africa, in S. obtusus Newell, 1971 and S. subobtusus
Newell, 1984 from off Chile, and in S. fuscus Viets,
1936, recorded from the Caribbean (Viets, 1936;
Newell, 1971, 1984; Bartsch, 1985b, 1994a; Otto,
2000a; Proches 2002).

In the just mentioned Australian species, as also in
S. latitarsus, the foveate ornamentation of the AE is
restricted to the lateral margins, ventrally the plate is
uniformly punctate. In contrast, in S. platyaspis the
AE is almost uniformly foveate. Females of the three
southern Pacific species, S. glareus, S. obtusus and S.
subobtusus, have a small PD and the ds-4 are situated
within the membraneous integument, whereas in S.
platyaspis that pair of setae is on the PD. Moreover, in
S. glareus and S. subobtusus the paired claws on tarsus
I are much more stout than in S. platyaspis, and
female S. obtusus have a shorter GA relative to the
size of GO.

Simognathus fuscus, a species from Bonaire (Viets,
1936), resembles S. platyaspis in shape and
ornamentation. Otto (2000a) used the length:width
ratio of the AD for discrimination, a quotient that
can be influenced by the mounting procedure. The
paratype of S. platyaspis, housed in the Zoological
Museum in Hamburg, Germany (ZMH A3/01), is
squeezed and the holotype (not seen by the author)
is suspected to be deformed, too. The length:width
ratio of the AD in the present specimen of S.
platyaspis is 1.65, that of the holotype of S. fuscus
1.69, hence almost the same. The rostrum of S.
fuscus is very short, about one fifth of the length of
the gnathosoma, but one fourth in S. platyaspis.

Distribution

Tropical Australia, Great Barrier Reef, 1-6 m,
coarse sand and coral rubble (Otto, 2000a) and
Dampier area (present record). Not bound to sandy
deposits.

Simognathus salebrosus sp. nov.

Figures 10A-J, 11A-F

Material Examined

Holotype

Male, Burrup Peninsula, east coast. King Bay,
20°38'S, 116°45'E, sandy tidal bank in a small river
emptying into King Bay, Western Australia,
Australia; 25 July 2000, coll. I. Bartsch (WAM
T52106).

Other Material Examined

Australia: Western Australia: 1 female, Dampier,
20°39'S, 116°42'E, sandy flat, lower part of tidal
slope, 1 August 2000, coll. I. Bartsch (WAM
T52107).

Etymology

Derived from Latin salebrosus, rough, uneven,
because of the rough surface of the plates.

Diagnosis

Idiosomal length 364-398 pm. AD and PD with
delicate punctation, their surface uniformly foveate.
AD longer than PD. OC elongate, covered by
membraneous integument. AE and GA fused.
Plates foveate apart from area representing anterior
part of GA. AE with large marginal epimeral fossae.
Area of epimera II and PE contiguous; lateral GA
and posterior part of PE almost articulating. Female
GA with four pairs of pgs. Male GA with three
pairs of outlying pgs and 26 pgs arranged in a ring
close around GO. Gnathosomal base dorsally and
marginally foveate; with triangular lamelliform
tectum. Palps short. Trochanter I large, with small
cylindrical base. Tibia I widest in basal third, then
evenly tapering. Posterior part of tibia I rotated
relative to axis of the leg. Tarsus I short, 0.17 of
length of tibia. Tarsi I to IV with 1, 1, 1, 1 ventral
setae and 2, 2, 1, 1 pas, respectively. Pectines of
claws of tarsi II to IV with numerous short tines.

Description

Male

Idiosoma. Male. Length 364 pm, width 189 pm.
AD and PD with intense punctation and superficial
foveae. Length of AD 167 pm, width 130 pm;
anterior margin undulate due to foveae (Figure
10A), posterior margin truncate. AD with ds-1 and
ds-3 as illustrated. OC reduced to short, elongate
sclerite which is covered by striae of membraneous
integument; its length 22 pm, width 7 pm. PD
slightly longer than AD, its length 175 pm, width
130 pm; anterior margin truncate. Pairs of ds-4 and
ds-5 slightly posterior to the level of insertions of
legs III and IV, respectively. Pair of ds-6 in marginal
position.

Ventral plates evenly punctate and foveate;
foveae absent posterior to camerostome, in an area
representing anterior part of GA and around GO

38

I. Bartsch

Figure 10 A-F. Simognathus salebrosus sp. nov. A, Idiosoma, dorsal, male; B, idiosoma, ventral, male; C, epimeral plate
with fossa and trochanter I, female; D, tectum, male; E, gnathosoma, ventral, male; F, genital area, male; G,
gnathosoma, lateral, female; H, leg I, medial and ventral, male; I, leg III, medial (trochanter in ventral
aspect), male; J, trochanter and basifemur III, medial, female. Scale = 50 pm (epf, epimeral fossa; epr,
epimeral process; tr, trochanter).

(Figure 10B). AE and GA fused to a ventral shield;
its length 289 pm, width 164 pm. Epimera II and PE
as well as posterior part of PE and lateral GA
contiguous. Epimeral processes I to IV forming
large lamellae. Anterior epimera with large
marginal fossae lateral to insertions of legs I (Figure
10C) and II. AE with three pairs of setae and rather
small epimeral vesicles. PE extending posteriad
beyond insertion of leg IV. GO small, length 45 pm,
width 27 pm, situated close to posterior margin of
idiosoma. GA with three pairs of outlying pgs and
26 pgs arranged in a ring close around GO. Genital
sclerites with three pairs of sgs (Figure 10F).

Gnathosoma. Length 115 pm, width 99 pm.
Gnathosomal base dorsally and marginally foveate
(Figure 10G), median part of ventral flank almost
smooth (Figure 10E). Dorsum with lamellar,
triangular tectum (Figure 10D). Basal pair of
maxillary setae long, distal pair short, slender. Palps
short, three-segmented. Second segment with
ventral seta but without protuberance; delicate
incision may represent fusion between P-2 and P-3.

Legs I to IV rather similar in length. Base of
trochanters I and II cylindrical, bent in an almost
right angle to longitudinal axis of the legs (Figure
10C). Trochanters III and IV elongate, almost as

Psammophilous halacarids from Western Australia

39

long as telofemora. Telofemora III and IV longer
than tibiae. Tibia I with cylindrical base, abruptly
expanding at 0.28 (relative to length of tibia), then
evenly tapering (Figure 10H). Posterior portion
rotated so that the ventral seta is in a medial
position. Tibiae II to IV claviform (Figures 101, 11B
and C). Tarsus I very short, hardly longer than high,
its median claw turned inward (Figure 11D and E ).
Tarsi II to IV at least three times longer than high.
Leg chaetotaxy (solenidia omitted, pas included):
leg I, 1, 2, 2, 4, 5, 6; leg II, 1, 2, 3, 4, 5, 6; legs III and
IV, 1, 1, 2, 3, 5, 5. Ventral spine of tibia I stout,
bluntly ending, without basal apophysis (Figure
11D); medial seta slender, flagelliform. Tibiae II to
IV with pair of wide, roughly bipectinate ventral
setae. Tarsi I to IV with three dorsal fossary setae;
on tarsus I dorsomedial fossary seta shorter than
dorsolateral one; on tarsi II to IV paired fossary
setae delicately serrate. Tarsi I to IV each with long
ventral seta, that seta on tarsus I basally wide, then
tapering, on tarsi II to IV ventral seta delicately
serrate. Tarsi I and II each with pair of pas, tarsi III
and IV with medial pas. Solenidion of tarsus I
setiform, famulus lamellar. Solenidion of tarsus II
setiform, situated on inside of medial fossa
membrane (Figure 11F).

Tarsus I with median claw and very slender
paired claws, these claws smooth. Paired claws of
tarsi II to IV large, with accessory process and
pectines; central sclerite minute, without distinct
claw-like process.

Female

Dorsal aspect resembling that of male. AE and
GA fused. GA with four pairs of pgs (Figure 11 A).

Remarks

Simognathus salebrosus is strikingly similar to S.
abnormalus Otto, 2000 and S. scutatus Bartsch, 1993,
species recorded from the Great Barrier Reef,
Queensland and Rottnest Island, Western Australia,
respectively (Otto, 2000a; Bartsch, 1993), the three
species have a ventral shield, short tarsus I and a
short palp which lacks a protuberance. S. scutatus is
easily distinguished from the others because all
ventral plates are fused, both in the female and
male, and its tibia I is almost equal in height for
most of its length. In contrast to males of S.
abnormalus, which bear two pairs of outlying setae,
the single male of S. salebrosus has three pairs of
such setae. The female GA and PE of S. salebrosus

Figure 11 A-F. Simognathus salebrosus sp. nov. A, Idiosoma, ventral, female; B, leg II, medial, female; C, leg IV, medial,
female; D, tibia and tarsus I, medial, female; E, posterior part of tibia and tarsus I, medial, female
(ventromedial seta and ventral spine of tibia omitted, lateral pas and claw omitted); F, tarsus II, medial,
female (lateral pas and claw omitted). Scale = 50 pm.

40

I. Bartsch

are almost contiguous but not fused. In S.
abnormalus variants are known with the GA and PE
either fused or separated (Otto, 2000a). Simogttathus
salebrosus and S. abnormalus differ in that in the
former species the AD is foveate throughout, in S.
abnormalus the anterolateral portions are non-
foveate.

With the shape of leg I, the tibia being highest in
the basal third, then tapering towards the rotated
distal portion, and a very short tarsus, this
Simognathus resembles Acaromantis species. The
three-segmented palps and the presence of paired
claws associated with the median claw are
characters of the genus Simognathus.

Conspicuous of S. salebrosus, and also of S.
abnormalus, is that epimera I and II form large
marginal fossae. The distal part of trochanters I and

II are within these epimeral fossae while the
articulation of the trochanters is protected by the
epimeral processes.

Distribution

Indian Ocean, Western Australia, Dampier. From
a tidal beach area.

Simognathus tener sp. nov.

Figures 12A-J, 13A-D

Material Examined

Holotype

Male, Dampier, 20°39'S, 116°42'E, Western
Australia, Australia; beach in recreation area, coarse

Figure 12 A-J. Simognathus tener sp. nov. A, Idiosoma, dorsal, male; B, idiosoma, ventral, male; C, genital area, male;

D, palp, medial, female; E, insertion of trochanter IV, ventral, male; F, gnathosoma, dorsal, male; G,
gnathosoma, lateral, female; H, gnathosoma, ventral, male; I, basifemur to tarsus I, medial, male; J, leg II,
medial, male. Scale = 50 pm.

Psammophilous halacarids from Western Australia

sand, mid-tide, 0-3 cm sediment depth, 3 August
2000, coll. I. Bartsch (WAM T52108).

Pamtype

Australia: Western Australia: 1 female, collecting
data same as above (WAM T52109).

Etymology

Specific name derived from tener, Latin, delicate,
referring to the delicate ornamentation of the plates.

Diagnosis

Idiosomal length 290-320 pm. Dorsal plates
punctate, surface with small, faint foveae. AD
shorter than PD. OC elongate, covered by
membraneous integument. Anterior margin of PD
ovate. AE and GA fused to elongate ventral shield;
its surface covered by very delicate foveae. Foveae
absent in area representing anterior portion of GA.
Gnathosomal base foveate. Dorsum with raised
cuticular markings, but without a dorsal lamella
extending beyond base of palps. Palps long, more
than half of idiosomal length. Second palpal
segment elongate, with ventral protuberance, seta
and an apical triangular lamella. Length of tibia I
twice its height. Ventral spine on tibia I long, evenly
tapering. Tarsi II to IV with six setae each,
parambulacral setae included (solenidia omitted).

Description

Male

Idiosoma. Male. Length 290 pm, width 150 pm;

41

length:width ratio 1.9. Dorsal plates evenly
punctate, surface with delicate foveae (Figure 12A).
Length of AD 130 pm, width 92 pm; posterior
margin truncate. Gland pores tiny, near lateral
margin. AD with ds-1 and ds-3, the latter removed
from lateral margin of plate. Pair of ds-2 on OC
which is hidden beneath striae of membraneous
integument. Length of PD 147 pm, width 102 pm;
anterior margin ovate. PD with ds-4 and ds-5 (in
holotype one of ds-4 lacking). Setae ds-6 in
marginal position.

Ventral plates evenly punctate; surface largely
covered by small, delicate foveae; these foveae
absent in an area representing anterior part of GA
and around GO (Figure 12B). AE and GA fused to a
ventral shield; its length 263 pm, width 122 pm.
Epimeral processes I to IV forming large lamellae
(Figures 12B and E). PE extending posteriad beyond
insertion of leg IV. AE with three pairs of setae; the
two posterior ones at almost the same level. GO
small, length 20 pm, width 12 pm. GA with one
pair of outlying setae and 31 slender setae around
GO. Spermatopositor extending beyond ring of pgs
(Figure 12C).

Gnathosoma. Coarsely foveate (Figures 12F-H); its
length 97 pm, width 75 pm. Dorsum of
gnathosomal base roughly textured, without
lamelliform tectum (Figure 12F); pair of lateral
lamella enclosing P-1 (Figure 12G). Palps slender;
second palpal segment long, about 0.6 of
gnathosomal length; segment with ventromedial
lamella and protuberance (Figure 12D), strong
ventral seta, and triangular, distolateral lamella.
The latter discernible in dorsolateral aspect.

Figure 13 A-D. Simognathus tener sp. nov. A, Idiosoma, ventral, female; B, leg III, medial, female; C, leg IV, medial,
female; D, tarsus I, lateral (medial setae and claw omitted, lateral pas in holotype broken), male. Scale = 50 pm.

42

I. Bartsch

Legs. Telofemur and tibia I almost equal in length.
Tibia II shorter than telofemur II. Base of tibia I
cylindrical, then abruptly widened and almost
equal in height in posterior half of segment (Figure
121). Tibiae II to IV clavate (Figures 12J, 13B and C).
Leg chaetotaxy (pas included, solenidion excluded):
leg I, 1, 2, 2, 4, 5, 6; leg II, 1, 2, 3, 4, 5, 6; legs III and
IV, 1, 1, 2, 3, 5, 6. Ventral spine of tibia I without
basal apophysis but evenly tapering; medial seta
flagellate. Tibiae II to IV each with two coarsely
bipectinate setae. Each of tarsi I to IV with three
dorsal setae, one ventral seta and a pair of pas. That
ventral seta on tarsus I basally stout, then tapering.
Solenidion of tarsus I setiform, famulus lamelliform
(Figure 13D).

Median claw on tarsus I large, smooth, paired
claws similar in length but slender, resembling
parambulacral setae. Paired claws of tarsi II to IV
with pectines, each with numerous tines.

Female

Length 320 pm. Dorsal aspect resembling that of
male. Ventral plates AE and GA fused (Figure 13A).
GA with four pairs of pgs.

Remarks

Simognathus tener and S. aspidiotus Otto, 2000
share numerous characters, the AE and GA form a
ventral shield; the males have a single pair of
outlying setae, the palps are long and slender, the
second segment is provided with a protuberance
and distal lamella; tarsi II to IV each bear one
ventral seta and a pair of pas. Simognathus tener has
more slender dorsal plates and more delicate foveae
than S. aspidiotus.

Species with a ventral shield are S. abnormalus, S.
aspidiotus Otto, 2000, S. clypeatus Otto, 2000, S.
gibberosus Bartsch, 1994, S. salebrosus, S. scutatus
Bartsch, 1993, S. tener, S. tropicalis Chatterjee & de
Troch, 2000, and S. uniscutatus Bartsch, 1994
(Bartsch 1993, 1994a; Chatterjee & de Troch, 2000;
Otto, 2000a).

In contrast to the other above species, the AD of
S. gibberosus bears a hyaline lens. S. abnormalus and
S. salebrosus have an Acaromantis- like tibia I,
elongate and rotated. In S. scutatus all ventral plates
are fused, there are no striae of membraneous
integument between AE and PE. Tarsi III and IV of
S. uniscutatus have a ventral seta and a single pas,
the medial but no lateral pas. In contrast, tarsi III
and IV of S. aspidiotus, S. clypeatus and S. tener each
bear a ventral seta and both a medial and lateral
pas; in S. clypeatus one of the pas is doubled. The
three species can be separated on the base of the
ornamentation of the dorsal and ventral plates; the
ventral plates are almost smooth in S. clypeatus,
delicately foveate in S. tener and coarsely foveate in
S. aspidiotus. The description of S. tropicalis, by
Chatterjee & de Troch (2000), presents no

information on the arrangement and number of the
pas.

According to Otto (2000a), the male of S.
gibberosus (housed in the WAM) has the AE and GA
separated. A re-examination of that male showed
that the AE and GA are fused to a ventral shield
but there is a rupture. Most likely, the slide had
been pressed seriously, a pressure and distortion
which caused a split in the integument of the mite
which was soaked and hardened by the mounting
medium (glycerin jelly). The rupture largely follows
the borders of the relevant plates. Hence, according
to present knowledge, males of S. gibberosus have a
ventral shield .

Distribution

Indian Ocean, Western Australia, Dampier. From
a tidal beach.

Simognathus uniscutatus Bartsch, 1994
Figure 14A-D

Simognathus uniscutatus Bartsch, 1994a: 145, figures
45-52.

Material Examined

Australia: Western Australia: 1 female, Enderby
Island, West of Rocky Head, 20°32.1'S, 116°26.7'E,
sponge garden, 2-13 m, unsorted sandy deposits, 3
August 2000, coll. C. Bryce (WAM T52110).

Diagnosis

Idiosomal length 390 pm. Dorsal plates densely
punctate and distinctly foveate. AD shorter than PD
(Figure 14A). Anterior margin of PD truncate. OC
reduced to oblong sclerite beneath striae of
integument. Pair of ds-2 in anterior edge of that
platelet. AE and GA fused (Figure 14B). This ventral
shield with small foveae except for a smooth median
portion representing anterior part of GA. AE with
large epimeral processes. Female GA with four pairs
of sgs. Gnathosomal base foveate; dorsally with
raised cuticular ornamentation but without tectum-
like lamella (Figure 14C). Second palpal segment
elongate, with ventral protuberance, a seta and a
triangular lamella. Leg chaetotaxy (solenidia
omitted, pas included): leg 1 , 1, 2, 2, 4, 5, 6; leg II, 1,
2, 3, 4, 5, 6; legs III and IV, 1, 1, 2, 3, 5, 5. Tibia I short,
with cylindrical base, then conspicuously widened
(Figure 14D); its ventral spine evenly tapering,
without apophysis. Pair of ventral spines on tibiae II
to IV distinctly bipectinate. Tarsi I and II each with a
wide ventral seta and a pair of eupathid pas; tarsi III
and IV with a ventral seta and a single medial pas.
On tarsus IV ventral seta and pas situated almost
adjacent. Tarsus I with large median and slender
paired claws. Paired claws of the following legs with
coarsely dentate pectines. Central sclerite with small
process.

Psammophilous halacarids from Western Australia

43

Figure 14 A-F. Simognathus uniscutatus Bartsch, 1994, female. A, Idiosoma, dorsal; B, idiosoma, ventral; C, part of
gnathosoma, dorsal; D, leg I, medial. Scale = 50 pm.

Remarks

The present female is larger than the holotype
female from Rottnest Island, Western Australia (390
pm vs 322 pm) and basifemur I bears two instead of
three setae, as present in the holotype of
Simognathus uniscutatus. Two setae on basifemur I is
the usual number in the majority of Simognathus
species and the presence of three setae is expected
to be an anomaly and of no taxonomic relevance.

Distribution

Western Australia, Rottnest Island and Dampier.

FAUNAL COMPARISON

From Australia, information on psammophilous
halacarid species is available from Rottnest Island
and Dampier, Western Australia, and the Great
Barrier Reef Marine Park and Coral Sea,
Queensland. Though the data sets are far from
complete, a preliminary comparison of the faunas
can be made.

Rottnest Island, at a latitude of about 32°S, is
characterized by numerous sandy beaches
separated by limestone platforms. The beaches,
with different exposure to waves and currents,
demonstrate a variety of grain size composition.
Rottnest Island lies in a transition zone between the
warm-temperate and tropical zone (Wells and
Walker, 1993).

Dampier is in the tropical arid zone, at about 20°S.
The coastline is dominated by muddy and silty flats
or mud-covered rocky platforms in the mid- and
lower tidal zone, rocks, boulders and mangroves in

the higher tidal area. Sandy deposits with coarse
and medium-sized grains are rare. The subtidal
sediment immediately off Dampier is rich in
organic material but around the outer islands of the
Dampier Archipelago there are coarse sediments.

The tropical Great Barrier Reef Marine Park and
adjacent Coral Sea cover a latitudinal area from
about 14° to 21 °S. There is a variety of different
substrata, amongst others areas with fine to coarse
sand and coral rubble.

The number and extent of collections performed
in the three areas Rottnest Island, Dampier and
Great Barrier Reef vary from sampling just once,
during a short stay of about two and a half weeks
(Rottnest Island, Dampier) to numerous collections
taken in a period of about two years (Great Barrier
Reef). Additional species (Table 1) are expected to
be found in all three areas.

Table 1 Halacarid genera and number of
psammophilous species recorded from the
Dampier area, Rottnest Island, the Great
Barrier Reef and Coral Sea.

Dampier

Rottnest

Island

Great Barrier Reef,
Coral Sea

Adacarus

2

2

4

Anomalohalacarus 1

1

0

Arhodeoporus

1

2

7

Copidognathus

1

7

14

Scaptognathides

2

1

3

Scaptognathus

1

2

8

Simognathus

4

7

12

44

I. Bartsch

Table 2 Psammophilous halacarid species from
Dampier area and notes on records from other
areas. +, record; no record

Dampier

Other areas

Actacarus pacificus

Actacarus festivus
Anomalolialacams dampierensis
Arhodeoporus corallicolus
Copidognathus meridianus
Scaptognathides hawaiiensis

Scaptognathides ornatus
Scaptognathus exquisitus
Simognathns salebrosus
Simognathus platyaspis
Simognathus tener
Simognathus uniscutatus

+ (Rottnest I., Hokkaido,
Hawaiian Is, Robinson
Crusoe I.)

+ (Great Barrier Reef)

+ (southern China,
Hawaiian Is)

+ (Hawaiian Is)

+ (Great Barrier Reef)

+ (Great Barrier Reef)

+ (Rottnest I.)

In the Dampier area, the collection from sandy
deposits and coral fragments contained 12 species,
seven of them have been recorded previously from
other areas, two species from the Indian and six
from the Pacific Ocean and adjacent basins, three
from the Great Barrier Reef and three from the
Hawaiian Islands (Table 2). The data on the
psammobiont halacarid fauna corroborates an
assumption of the presence of a rather uniform
tropical coastal fauna from North West Cape across
the northern Australian coastline to Queensland
and also to the Hawaiian Islands. Yet, because of
the small size of psammobiont halacarids and hence
difficulties in recognizing details, the presence of
cryptic species cannot be excluded. Studies on the
tidal rhombognathine halacarids from Dampier also
demonstrated close similarities with the fauna of
the northeastern Australia (Bartsch, 2003a).

ACKNOWLEDGEMENTS

The samples were taken in July and August 2000
during the Woodside Dampier Marine Biological
Workshop which was organized by Dr Fred Wells
and Di Jones, Western Australian Museum, and
Prof. Diana Walker, The University of Western
Australia. Clay Bryce provided me with subtidal
sediments from the outer islands. The workshop
was sponsored by Woodside Energy Ltd and
accomodation was provided by Hamersley Iron
Pty. Ltd. My flight to Australia was financed by the
German Research Foundation (DFG). Dr H. Dastych
arranged loan of specimens housed in the
Zoological Institute and Zoological Museum in
Hamburg. To all my sincerest thanks.

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Wagler (eds), Die Tierwelt der Nord- und Ostsee. la: 1-
238,

Viets, K. (1936). Zoologische Ergebnisse einer Reise nach
Bonaire, Curasao und Aruba im Jahre 1930. No. 18.
Halacariden aus Westindien. Zoologische Jahrbiicher fiir
Systematik, Okologie und Tiergeographie 67: 389-424.

Wells, F.E. and Walker, D.l. (1993). Introduction. In F.E.
Wells, D.L Walker, H. Kirkman and R. Lethbridge
(eds), The Marine Flora and Fauna of Rottnest Island,
Western Australia:l-10. Western Australian Museum,
Perth.

Manuscript received 10 January 2003; accepted 4 April 2003

Records of the Western Australian Museum 22: 47-65 (2003).

Taxonomic status of the ricefield rat Rattus argentiventer (Robinson
and Kloss, 1916) (Rodentia) from Thailand, Malaysia and
Indonesia based on morphological variation

Ibnu Maryanto

Museum Zoologicum Bogoriense, R&D Centre For Biology,

Indonesia Institute of Sciences (LIPI). Jl. Raya Jakarta Bogor Km 46, Cibinong, Bogor, Indonesa
e-mail: ibnu_mar@yahoo.com; mzb@indo.net. id

Abstract - Comparison of cranial and external characters of 212 ricefield rats
Rattus argentiventer (Robinson and Kloss 1916), by univariate and multivariate
statistical analyses, was carried out on specimens from Thailand, Malaysia,
and Indonesia (Sumatra, Kalimantan, Java, Bali, Lesser Sunda, Sulawesi and
Irian Jaya). These comparisons indicate that ricefield rats from Thailand to
Irian Jaya fall into four groups: a Java group (Thailand, Malaysia, Sumatra
and Java), a Bali-Sulawesi group (Bali, Sulawesi, Lombok, Sumbawa,
Adonara, Sangeang, Rinca, Flores, Lembata, Alor, Timor and Tanimbar
Islands); a Sumba group (Sumba Island) and a Kalimantan group (Kalimantan
Island). The specimen from Irian Jaya was placed in the Java group. The
taxon from Kalimantan is described as a new subspecies, R. a. kalimantan.

The multiple regression analysis indicated that while skull, dentary, dental,
and external characters of the ricefield rat do not exhibit sexual dimorphism,
age or interactions between islands, sex and /or age does influence some
characters.

Key words: Ricefield rat, Rattus argentiventer, morphological variation.

Thailand, Malaysia, Indonesia.

INTRODUCTION

Recent reappraisal of the mammalian fauna of the
Lesser Sunda and Sunda Islands has led to a
dramatically different view of the biogeographic
boundaries in the Oriental and Australian regions
from the traditional ones based on Wallace's and
Weber's lines (Kichener et al., 1990; Kitchener and
Maryanto, 1993; Kitchener et al., 1995). The
Indonesian archipelago presents unique
opportunities to study morphological variation.
Comprehensive re-evaluation of morphology in
many cases resulted in the discovery of new species
and subspecies. In Indonesia such re-evaluation has
been conducted, for example, in bats, rodents and
primates (Kitchener and Maryanto, 1993; Maryanto
et al, 1997; Maryanto and Kitchener, 2000).

The ricefield rat, Rattus argentiventer, a rodent
species found in ricefields, is often a major pest. It is
distributed through Thailand, Vietnam, Cambodia,
Laos, Malaysia, Sumatra, Java, Kalimantan,
Sulawesi, the Philippines, Lesser Sunda, and has
been introduced to New Guinea (van Strien, 1986;
Flannery, 1990; Suyanto et al, 1999). There are
indications that R. argentiventer has different
behaviors in different locations (Maryanto, 1991).
Recent authors recognized one species (Musser,
1977; van Strein, 1986), however Kitchener and

Suyanto (1996) argued that a detailed examination
of morphological variation between island
populations is needed to resolve the taxonomic
situation of widespread species such as the ricefield
rat, especially in the Indo-Australian region.

A number of recent authors have discussed the
taxonomic status of the ricefield rat. Prior to
Musser's classification of species limits in Asian
Rattus, most authors treated R. argentiventer as a
subspecies of Rattus rattus (Olerman, 1951; Schwarz
and Schwarz, 1967). Musser (1972) initially treated
specimens of R. argentiventer from Bali, Lombok,
Sumbawa, Flores and Timor as a subspecies of R.
rattus, classifying them as Rattus rattus bali. He
further considered that R. r. satumus from Sumba
Island was synonymous with R. r. bali. Specimens
from Java were first described as R. r. brevicaudatus
(Horst and Raadt, 1918) however Chasen (1940)
considered this as a synonym of R. r. argentiventer.
Specimens from Sulawesi described as R. pesticulus
Thomas, 1921 were referred by Musser (1977) to R.
argentiventer. Kitchener et al (1990) referred
specimens from Lombok to R. a. bali.

This paper reports on a taxonomic reappraisal of
R. argentiventer, as defined above, from Thailand,
Malaysia, and Indonesia, based on an examination
of morphological variation. Specimens from

48

I. Maryanto

mainland Southeast Asia north of the Isthmus of
Kra and from the Phillipine islands have not been
examined as part of this study.

MATERIALS AND METHODS

A total of 212 adult specimens (see Appendix 1)
were studied. The specimens were collected from
Thailand, Malaysia, Sumatra, Kalimantan, Java,
Sulawesi, Bali, Lombok, Sumbawa, Sangeang,
Flores, Adonara, Lembata, Alor, Sumba, Timor,
Tanimbar, and from Irian Jaya (Figure 1). All
specimens studied are in the collection of Museum
Zoologicum Bogoriense (MZB), the Western
Australian Museum (WAM) and the Raffles
Museum, Zoological Reference Collection,
Singapore (ZRC). A specimen was judged to be
adult if both the basioccipital/basisphenoid and
basisphenoid/presphenoid sutures were fused.
Twenty-five measurements of skull, dental and
dentary characters and four external characters
were measured with calipers to 0.01 mm. The skull
characters used were: BB, bulla breadth; BH, bulla
height; BL, bulla length; BOZP, breadth of
zygomatic plate; BS, braincase breadth; CBL,
condillo basal length; GSL, greatest skull length;
HB, braincase height; IFB, incisive foramen breadth;
IL, incisive foramen length; IO, interorbital breadth;
LOD, length of diastema; LOP, length of palatal;
M 3 W, upper molar 1 width; M'M 1 , upper molar 1 to
molar 1; M 2 M 2 , upper molar 2 to molar 2.; M 3 M 3 ,
upper molar 3 to molar 3; MSF, mesopterygoid

fossa width; NB, nasal width; NL, nasal length; DL,
dentary length; POW, post orbital width; RAP,
ramus angular process; TR, upper tooth row; ZB,
zygomatic width. The external characters were
HBL, head and body length; TL, tail length; E, ear
length and HF, hind foot length. The complete
character measurements are shown in Figure 2. Tine
pelage descriptions follow the color terminology of
Konerup and Wanscher (1983).

The statistical analyses were run in two steps,
univariate analysis and multivariate analysis, as
described in Maryanto and Sinaga (1998). Sexes
were analyzed separately, as were the skull and
external characters. Discriminant Function Analysis
(DFA) was initially run for all characters and with
all islands as separate groups for island groups.
Islands with fewer than four specimens were
treated as unallocated island groups. The DFA was
then run for a subset of five or more skull
characters, based on the criteria of minimized
Wilk's lambda to select the best discriminating
variables results obtained with the reduced
character set were very similar to those based on all
characters (Kitchener et al. 1993)

RESULTS AND DISCUSSION
Univariate statistics

Mean, standard deviation, minimum and
maximum values and sample size are presented in
Table 1. These suggest that, except for IFB, MSF,

Figure 1 Localities of Rattus argentiventer specimens used in this study.

Taxonomic status of the ricefield rat, Rattus argentiventer

49

M 2 M 2 and M 3 M 3 , all skull characters in males were
slightly larger than in females. Furthermore, the
averages of males and females from each group
showed that the characters showing sexual
differences differed in different groups. For
example: The characters in males from Java were
slightly larger than in females; nearly all characters
in males from the Bali-Sulawesi group were larger
than in females, except for IO, MSF, and fvTM 1 -

M 3 M 3 . Furthermore nearly all characters in males
from Sumba were smaller than in females except
for GSL, NB, and IFB; and lastly nearly all
characters in males from Kalimantan were smaller
than in females except for E, HF, MSF, BH, and
M*W (Table 1)

Multiple regression

Multiple regression analysis was employed to

50

I. Maryanto

DF 1

DF 1

DF 2

Figure 3 Plot of the female Canonical variate on the following island group populations; ♦ Java group (Thailand,
Malaysia, Sumatra, and Java); ■ Kalimantan group (Kalimantan Island); O Bali group (Bali, Lombok,
Sumbawa, Sangeang, Komodo, Flores, Adonara, Lembata, Alor, Timor and Tanimbar) and * Sumba group
(Sumba Island). A. Discriminant Function (DF) 1 vs. Discriminant Function 2 . B. Discriminant Function 1 vs.
Discriminant Function 3 and C. Function 2 vs. Function 3.

Taxonomic status of the ricefield rat, Rattus argentiventer

Table 1 Male and female measurements of the skull, dentary and external characters (in mm).

51

Group SEX LB TL E HF

Female N

Mean

Female N

Mean

Std. Deviation
Minimum

N

Mean

Std. Deviation

Minimum

Maximum

Std. Deviation
Minimum
Maximum
N

Mean

Std. Deviation

Minimum

Maximum

N

Mean

Std. Deviation

Minimum

Maximum

Java Male

Total

Sumba Male

Total

Bali Male

Total

Kalimantan Male

Maximum

N

Mean

Std. Deviation

Minimum

Maximum

N

Mean

Std. Deviation

Minimum

Maximum

Maximum
N

Mean
Std. Deviation
Minimum
Maximum

N

Mean

Std. Deviation
Minimum
Maximum
N

Mean

Std. Deviation
Minimum
Maximum
N

Mean

Std. Deviation

Minimum

Maximum

39 43

182.82 168.19

16.99 13.13

154.00 120.00

240.00 192.00

39 43

172.38 155.07

10.56 15.95

147.00 119.00

199.00 190.00

78 86

177.60 161.63

15.00 15.95

147.00 119.00

240.00 192.00

2 2

152.00 151.50

25.46 27.58

134.00 132.00

170.00 171.00

8 7

180.81 168.43

20.40 20.39

148.00 145.00

200.00 191.00

10 9

175.05 164.67

23.31 21.51

134.00 132.00

200.00 191.00

21 21

175.98 170.60

12.57 13.24

157.00 144.00

213.00 201.00

22 22

173.84 161.77

14.12 17.51

145.00 120.00

201.00 194.00

43 43

174.88 166.08

13.27 16.02

145.00 120.00

213.00 201.00

5 5

167.50 148.10

18.43 10.85

139.50 135.50

187.00 163.00

11 11

174.91 156.32

14.17 11.09

150.00 136.00

199.00 172.00

16 16

172.59 153.75

15.39 11.35

139.50 135.50

199.00 172.00

43 43

20.05 34.72

2.09 2.75

16.00 27.00

28.00 39.00

43 43

19.34 33.33

1.68 3.04

16.00 29.00

23.00 43.00

86 86

19.69 34.02

1.92 2.96

16.00 27.00

28.00 43.00

2 2

18.00 32.50

1.41 0.00

17.00 32.50

19.00 32.50

8 8

21.06 34.00

2.21 1.77

17.00 31.00

24.00 36.00

10 10

20.45 33.70

2.39 1.69

17.00 31.00

24.00 36.00

20 20

20.40 34.13

1.70 2.17

17.00 29.00

23.00 37.00

22 22

20.80 33.82

1.84 2.14

17.00 30.50

24.00 39.00

42 42

20.61 33.96

1.76 2.13

17.00 29.00

24.00 39.00

5 5

20.30 35.00

0.84 2.83

19.00 32.00

21.00 38.00

10 11

19.50 34.91

1.18 2.59

17.00 30.00

21.00 38.00

15 16

19.77 34.94

1.12 2.57

17.00 30.00

21.00 38.00

Female

Total

Female N

Mean

Std. Deviation
Minimum

GSL ZB HB

52

53

53

41.02

19.74

11.76

1.66

0.88

0.63

37.50

17.80

10.80

44.50

21.40

15.00

52

53

53

39.79

19.53

11.59

1.48

0.74

0.38

37.20

18.00

10.80

44.00

21.50

12.40

104

106

106

40.40

19.64

11.68

1.68

0.82

0.53

37.20

17.80

10.80

44.50

21.50

15.00

5

6

6

41.76

19.51

11.64

3.91

1.43

0.37

37.60

18.20

11.30

46.70

21.90

12.20

9

9

9

41.68

20.39

12.22

2.80

1.21

0.74

38.80

18.40

11.00

47.00

22.00

13.10

14

15

15

41.71

20.04

11.99

3.09

1.33

0.67

37.60

18.20

11.00

47.00

22.00

13.10

36

35

35

41.44

19.83

11.88

2.70

1.30

0.69

36.60

17.10

10.50

46.50

22.70

13.50

37

37

36

40.73

19.76

11.78

2.27

1.42

0.64

36.50

17.30

10.50

45.60

23.20

13.20

73

72

71

41.08

19.79

11.83

2.50

1.35

0.66

36.50

17.10

10.50

46.50

23.20

13.50

5

5

5

40.17

19.11

11.60

1.68

1.04

0.35

38.10

18.20

11.30

42.50

20.70

12.10

12

12

12

40.99

19.90

11.78

1.52

0.76

0.42

38.90

18.70

10.80

44.50

21.00

12.30

17

17

17

40.75

19.67

11.73

1.57

0.89

0.40

38.10

18.20

10.80

44.50

21.00

12.30

NL NB RAP

52

53

51

14.48

4.41

13.01

0.99

0.25

0.96

12.30

4.00

11.30

16.80

5.20

16.90

52

53

53

14.01

4.33

12.67

0.79

0.30

0.79

12.30

3.80

11.30

15.40

5.00

14.90

104

106

104

14.24

4.37

12.84

0.92

0.28

0.89

12.30

3.80

11.30

16.80

5.20

16.90

6

6

5

15.03

4.99

12.05

2.02

0.45

1.13

12.70

4.70

10.90

18.40

5.70

13.90

9

9

8

15.11

4.83

13.40

1.32

0.20

1.28

13.70

4.50

11.60

17.90

5.10

15.20

15

15

13

15.08

4.89

12.88

1.57

0.32

1.36

12.70

4.50

10.90

18.40

5.70

15.20

35

36

34

14.93

4.79

13.05

1.27

0.45

2.07

12.20

4.00

10.70

17.50

5.90

22.50

37

37

36

14.58

4.68

12.79

1.00

0.44

2.01

12.40

4.00

10.50

16.70

5.60

22.70

72

73

70

14.75

4.74

12.92

1.15

0.44

2.03

12.20

4.00

10.50

17.50

5.90

22.70

5

5

5

13.86

4.87

12.85

1.06

0.35

0.86

12.30

4.60

12.00

15.10

5.40

14.10

12

12

12

14.67

4.90

13.20

0.76

0.24

0.62

13.00

4.60

12.30

15.80

5.20

14.30

17

17

17

14.43

4.89

13.09

0.91

0.27

0.69

12.30

4.60

12.00

15.80

5.40

14.30

52

Table 1 (cont.)

I. Maryanto

Group

SEX

LB

TL

E

HF

GSL

ZB

HB

NL

NB

RAP

Average

- All combined

Male

N

67

71

70

70

98

99

99

98

100

95

Mean

178.61

167.01

20.11

34.51

41.17

19.73

11.79

14.64

4.60

12.97

Std. Deviation

16.97

14.43

1.92

2.56

2.22

1.08

0.63

1.19

0.41

1.46

Minimum

134.00

120.00

16.00

27.00

36.60

17.10

10.50

12.20

4.00

10.70

Maximum

240.00

201.00

28.00

39.00

46.70

22.70

15.00

18.40

5.90

22.50

Female

N

80

83

83

84

110

111

110

110

111

109

Mean

173.98

158.14

19.91

33.73

40.39

19.71

11.72

14.36

4.55

12.82

Std. Deviation

13.23

16.50

1.85

2.68

1.98

1.07

0.54

0.97

0.40

1.34

Minimum

145.00

119.00

16.00

29.00

36.50

17.30

10.50

12.30

3.80

10.50

Maximum

201.00

194.00

24.00

43.00

47.00

23.20

13.20

17.90

5.60

22.70

Total

N

147

154

153

154

208

210

209

208

211

204

Mean

176.09

162.23

20.00

34.08

40.75

19.72

11.75

14.49

4.57

12.89

Std. Deviation

15.18

16.15

1.88

2.65

2.13

1.07

0.58

1.09

0.41

1.40

Minimum

134.00

119.00

16.00

27.00

36.50

17.10

10.50

12.20

3.80

10.50

Maximum

240.00

201.00

28.00

43.00

47.00

23.20

15.00

18.40

5.90

22.70

Group

SEX

DL

M’W

M*L

IO

BOZP

LOP

TR

IL

MSF

IFB

Java

Male

N

52

52

51

53

53

53

53

53

53

53

Mean

22.80

2.10

3.19

5.71

4.87

19.14

7.00

7.68

2.36

2.59

Std. Deviation

1.04

0.11

0.22

0.26

0.39

0.92

0.32

0.48

0.22

0.24

Minimum

20.70

1.90

2.30

5.10

4.00

17.60

5.70

6.60

1.90

2.20

Maximum

24.60

2.30

3.50

6.40

5.90

21.40

7.50

8.50

2.90

3.20

Female

N

53

53

51

53

53

53

53

53

53

53

Mean

22.47

2.08

3.20

5.55

4.87

18.89

6.87

7.44

2.36

2.55

Std. Deviation

1.26

0.09

0.20

0.23

0.41

0.89

0.27

0.81

0.20

0.25

Minimum

20.50

1.90

2.80

5.00

4.00

17.10

6.30

2.80

1.90

2.10

Maximum

26.00

2.20

3.70

6.00

6.00

21.40

7.40

8.70

2.80

3.30

Total

N

105

105

102

106

106

106

106

106

106

106

Mean

22.63

2.09

3.19

5.63

4.87

19.01

6.94

7.56

2.36

2.57

Std. Deviation

1.16

0.10

0.21

0.26

0.40

0.91

0.30

0.67

0.21

0.25

Minimum

20.50

1.90

2.30

5.00

4.00

17.10

5.70

2.80

1.90

2.10

Maximum

26.00

2.30

3.70

6.40

6.00

21.40

7.50

8.70

2.90

3.30

Sumba

Male

N

6

6

2

6

6

6

6

6

6

6

Mean

23.11

2.19

3.21

5.70

4.71

19.23

7.07

7.83

2.43

2.60

Std. Deviation

1.86

0.06

0.08

0.39

0.45

1.48

0.09

0.58

0.14

0.38

Minimum

21.50

2.10

3.20

5.30

4.30

18.00

7.00

7.40

2.30

2.30

Maximum

25.80

2.30

3.30

6.40

5.60

21.80

7.20

8.70

2.60

3.20

Female

N

9

9

9

9

9

9

9

9

9

9

Mean

23.78

2.23

3.36

5.79

4.89

20.29

7.37

8.00

2.59

2.52

Std. Deviation

1.76

0.12

0.22

0.29

0.51

1.43

0.22

0.57

0.20

0.22

Minimum

21.90

2.00

2.80

5.40

4.40

18.70

7.10

7.30

2.20

2.10

Maximum

26.20

2.40

3.60

6.20

6.00

22.80

7.80

9.10

2.90

2.90

Total

N

15

15

11

15

15

15

15

15

15

15

Mean

23.51

2.21

3.33

5.75

4.82

19.86

7.25

7.93

2.53

2.55

Std. Deviation

1.77

0.10

0.21

0.32

0.48

1.50

0.23

0.56

0.19

0.29

Minimum

21.50

2.00

2.80

5.30

4.30

18.00

7.00

7.30

2.20

2.10

Maximum

26.20

2.40

3.60

6.40

6.00

22.80

7.80

9.10

2.90

3.20

Bali

Male

N

36

36

23

36

36

34

36

36

35

34

Mean

22.60

2.14

3.30

6.06

4.78

19.80

7.10

7.85

2.33

2.58

Std. Deviation

2.32

0.16

0.25

0.29

0.45

1.61

0.38

0.80

0.24

0.26

Minimum

13.40

1.70

2.80

5.60

3.70

17.00

6.40

6.60

1.90

2.20

Maximum

27.10

2.40

3.80

6.80

5.70

24.00

7.80

9.70

2.90

3.00

Female

N

37

37

20

37

37

37

37

37

37

37

Mean

22.62

2.13

3.25

6.09

4.64

19.64

7.15

7.41

2.40

2.58

Std. Deviation

2.37

0.13

0.25

0.40

0.46

1.36

0.37

0.63

0.30

0.28

Minimum

12.50

1.70

2.80

5.30

3.70

17.60

6.40

6.00

1.90

2.20

Maximum

26.60

2.40

3.80

6.90

5.70

23.20

8.00

9.50

3.10

3.20

Total

N

73

73

43

73

73

71

73

73

72

71

Mean

22.61

2.14

3.27

6.07

4.71

19.72

7.13

7.63

2.36

2.58

Std. Deviation

2.33

0.14

0.25

0.35

0.46

1.48

0.37

0.74

0.28

0.27

Minimum

12.50

1.70

2.80

5.30

3.70

17.00

6.40

6.00

1.90

2.20

Maximum

27.10

2.40

3.80

6.90

5.70

24.00

8.00

9.70

3.10

3.20

Taxonomic status of the ricefield rat, Rattus argentiventer

53

Group

SEX

DL

M'W

M'L

IO

BOZP

LOP

TR

IL

MSF

IFB

Kalimantan

Male

N

5

5

1

5

5

5

5

5

5

5

Mean

22.75

2.20

3.23

5.61

4.61

19.41

6.94

7.45

2.35

2.47

Std. Deviation

0.76

0.08

0.20

0.33

1.00

0.10

0.45

0.12

0.28

Minimum

22.10

2.10

3.20

5.40

4.20

18.30

6.80

6.80

2.30

2.10

Maximum

24.00

2.30

3.20

5.80

5.00

21.00

7.00

8.00

2.60

2.90

Female

N

12

12

4

12

12

12

12

12

12

12

Mean

23.54

2.19

3.28

5.85

4.90

19.68

7.08

7.48

2.29

2.59

Std. Deviation

1.04

0.10

0.12

0.20

0.33

0.75

0.18

0.44

0.25

0.21

Minimum

22.10

2.00

3.10

5.60

4.20

18.80

6.80

6.60

2.00

2.20

Maximum

25.60

2.30

3.40

6.20

5.40

21.20

7.50

8.00

2.90

3.00

Total

N

17

17

5

17

17

17

17

17

17

17

Mean

23.31

2.19

3.27

5.78

4.81

19.60

7.04

7.47

2.31

2.56

Std. Deviation

1.02

0.10

0.11

0.22

0.35

0.81

0.17

0.43

0.22

0.23

Minimum

22.10

2.00

3.10

5.40

4.20

18.30

6.80

6.60

2.00

2.10

Maximum

25.60

2.30

3.40

6.20

5.40

21.20

7.50

8.00

2.90

3.00

Average - All combined

Male

N

99

99

77

100

100

98

100

100

99

98

Mean

22.74

2.12

3.22

5.83

4.82

19.39

7.04

7.74

2.35

2.58

Std. Deviation

1.64

0.13

0.23

0.32

0.41

1.26

0.33

0.62

0.22

0.26

Minimum

13.40

1.70

2.30

5.10

3.70

17.00

5.70

6.60

1.90

2.10

Maximum

27.10

2.40

3.80

6.80

5.90

24.00

7.80

9.70

2.90

3.20

Female

N

111

111

84

111

111

111

111

111

111

111

Mean

22.74

2.12

3.23

5.78

4.80

19.34

7.03

7.48

2.38

2.56

Std. Deviation

1.77

0.12

0.21

0.38

0.44

1.18

0.33

0.71

0.25

0.25

Minimum

12.50

1.70

2.80

5.00

3.70

17.10

6.30

2.80

1.90

2.10

Maximum

26.60

2.40

3.80

6.90

6.00

23.20

8.00

9.50

3.10

3.30

Total

N

210

210

161

211

211

209

211

211

210

209

Mean

22.74

2.12

3.23

5.80

4.81

19.36

7.03

7.60

2.37

2.57

Std. Deviation

1.70

0.13

0.22

0.35

0.43

1.22

0.33

0.68

0.24

0.25

Minimum

12.50

1.70

2.30

5.00

3.70

17.00

5.70

2.80

1.90

2.10

Maximum

27.10

2.40

3.80

6.90

6.00

24.00

8.00

9.70

3.10

3.30

Group

SEX

BL

BH

BB

LOD

CBL

M'M 1

M 2 M 2

M 3 M 3

BS

POW

Java

Male

N

53

53

53

53

51

51

50

50

51

51

Mean

7.55

7.07

4.19

11.85

40.24

3.93

4.31

4.74

15.74

16.34

Std. Deviation

0.38

0.30

0.33

0.72

2.69

0.35

0.37

0.35

0.57

0.54

Minimum

6.50

6.20

3.50

9.90

26.30

3.14

3.50

4.03

14.58

15.27

Maximum

8.30

7.60

5.30

13.00

44.60

4.58

5.04

5.62

16.97

17.42

Female

N

53

52

53

53

52

53

53

53

53

53

Mean

7.38

6.92

4.10

11.52

39.22

3.85

4.17

4.68

15.53

16.26

Std. Deviation

0.33

0.24

0.27

0.56

1.45

0.42

0.41

0.39

0.61

0.62

Minimum

6.80

6.40

3.60

9.90

36.80

2.98

3.41

3.97

13.71

14.89

Maximum

8.10

7.40

4.80

12.80

43.00

4.78

5.34

5.73

16.89

17.92

Total

N

106

105

106

106

103

104

103

103

104

104

Mean

7.46

6.99

4.15

11.68

39.72

3.89

4.24

4.71

15.64

16.30

Std. Deviation

0.36

0.28

0.30

0.66

2.21

0.39

0.39

0.37

0.60

0.58

Minimum

6.50

6.20

3.50

9.90

26.30

2.98

3.41

3.97

13.71

14.89

Maximum

8.30

7.60

5.30

13.00

44.60

4.78

5.34

5.73

16.97

17.92

Sumba

Male

N

6

6

6

6

6

6

6

6

6

6

Mean

7.71

7.21

4.41

11.79

41.07

3.61

3.93

4.35

16.14

16.33

Std. Deviation

0.52

0.39

0.49

1.17

3.67

0.45

0.52

0.51

0.86

0.39

Minimum

7.00

6.80

3.90

11.10

36.90

3.01

3.24

3.75

14.91

15.81

Maximum

8.60

7.90

5.20

14.20

46.60

4.05

4.54

4.89

17.19

16.90

Female

N

9

9

9

9

9

9

8

8

9

9

Mean

8.08

7.48

4.48

12.65

41.42

4.02

4.44

4.86

16.52

16.79

Std. Deviation

0.68

0.41

0.21

1.00

3.30

0.49

0.59

0.57

0.83

0.63

Minimum

7.30

6.90

4.00

11.50

37.70

3.27

3.39

3.83

15.53

15.98

Maximum

9.00

8.00

4.80

14.10

47.50

4.68

5.24

5.56

18.18

17.71

Total

N

15

15

15

15

15

15

14

14

15

15

Mean

7.93

7.37

4.45

12.31

41.28

3.86

4.22

4.64

16.37

16.61

Std. Deviation

0.63

0.41

0.34

1.12

3.33

0.50

0.60

0.59

0.83

0.58

Minimum

7.00

6.80

3.90

11.10

36.90

3.01

3.24

3.75

14.91

15.81

Maximum

9.00

8.00

5.20

14.20

47.50

4.68

5.24

5.56

18.18

17.71

54

I. Maryanto

Table 1 (cont.)

Group SEX BL BH BB LOD CBL M'M 1 M 2 M 2 M 3 M 3 BS POW

Bali Male

N

35

35

35

36

34

29

29

29

31

31

Mean

7.49

7.00

4.19

12.14

40.67

3.92

4.29

4.73

15.98

16.67

Std. Deviation

0.39

0.36

0.28

1.23

3.39

0.45

0.46

0.57

0.69

0.67

Minimum

6.70

6.10

3.70

10.10

28.10

3.06

3.53

3.82

14.53

15.36

Maximum

8.40

7.60

4.70

15.40

46.10

4.70

5.20

5.64

17.21

18.90

Female

N

37

37

37

37

37

35

35

35

35

34

Mean

7.43

6.95

4.14

11.80

40.13

3.97

4.47

4.97

15.70

16.52

Std. Deviation

0.43

0.33

0.26

1.05

2.42

0.53

0.60

0.58

0.84

0.59

Minimum

6.40

6.10

3.70

9.50

35.60

3.10

3.39

4.11

14.09

15.44

Maximum

8.30

7.60

4.60

15.00

45.30

5.01

5.85

6.15

17.32

17.67

Total

N

72

72

72

73

71

64

64

64

66

65

Mean

7.46

6.97

4.17

11.97

40.39

3.95

4.39

4.86

15.83

16.59

Std. Deviation

0.41

0.34

0.27

1.15

2.92

0.49

0.54

0.58

0.78

0.63

Minimum

6.40

6.10

3.70

9.50

28.10

3.06

3.39

3.82

14.09

15.36

Maximum

8.40

7.60

4.70

15.40

46.10

5.01

5.85

6.15

17.32

18.90

Kalimantan Male

N

5

5

5

5

5

5

5

5

5

5

Mean

7.57

7.02

4.27

11.64

39.39

3.97

4.22

4.73

16.34

15.92

Std. Deviation

0.32

0.25

0.40

0.45

1.82

0.22

0.19

0.23

0.61

0.38

Minimum

7.10

6.60

3.90

11.10

37.10

3.74

3.89

4.36

15.40

15.51

Maximum

7.90

7.20

4.90

12.10

42.00

4.25

4.34

4.95

17.10

16.54

Female

N

12

12

12

12

12

12

12

12

12

12

Mean

7.57

7.01

4.34

11.89

40.41

3.99

4.30

4.74

16.34

16.12

Std. Deviation

0.28

0.21

0.36

0.45

1.49

0.23

0.34

0.43

0.78

0.44

Minimum

7.30

6.70

3.60

11.40

38.70

3.59

3.66

4.17

14.85

14.97

Maximum

8.20

7.30

4.70

13.00

43.60

4.38

4.88

5.42

17.39

16.68

Total

N

17

17

17

17

17

17

17

17

17

17

Mean

7.57

7.01

4.32

11.82

40.11

3.98

4.27

4.74

16.34

16.06

Std. Deviation

0.28

0.21

0.36

0.46

1.61

0.22

0.30

0.37

0.72

0.42

Minimum

7.10

6.60

3.60

11.10

37.10

3.59

3.66

4.17

14.85

14.97

Maximum

8.20

7.30

4.90

13.00

43.60

4.38

4.88

5.42

17.39

16.68

Average - All combined

Male

N

99

99

99

100

96

91

90

90

93

93

Mean

7.54

7.05

4.21

11.94

40.40

3.91

4.27

4.71

15.88

16.43

Std. Deviation

0.39

0.33

0.33

0.96

2.96

0.39

0.41

0.44

0.65

0.60

Minimum

6.50

6.10

3.50

9.90

26.30

3.01

3.24

3.75

14.53

15.27

Maximum

8.60

7.90

5.30

15.40

46.60

4.70

5.20

5.64

17.21

18.90

Female

N

111

110

111

111

110

109

108

108

109

108

Mean

7.47

6.98

4.17

11.74

39.83

3.91

4.30

4.79

15.76

16.37

Std. Deviation

0.44

0.32

0.29

0.84

2.10

0.44

0.50

0.49

0.79

0.62

Minimum

6.40

6.10

3.60

9.50

35.60

2.98

3.39

3.33

13.71

14.89

Maximum

9.00

8.00

4.80

15.00

47.50

5.01

5.85

6.15

18.18

17.92

Total

N

210

209

210

211

206

200

198

198

202

201

Mean

7.50

7.02

4.19

11.84

40.10

3.91

4.29

4.76

15.81

16.40

Std. Deviation

0.41

0.32

0.31

0.90

2.55

0.42

0.46

0.47

0.73

0.61

Minimum

6.40

6.10

3.50

9.50

26.30

2.98

3.24

3.75

13.71

14.89

Maximum

9.00

8.00

5.30

15.40

47.50

5.01

5.85

6.15

18.18

18.90

determine the influences of sex, age, localities and
the interactions between these factors. All localities
were included in the analysis (Malaysia, Sumatra,
Java, Kalimantan, Sulawesi, Bali, Lombok,
Sumbawa, Sangeang, Flores, Sumba, Adonara,
Lembata, Alor, Timor, Tanimbar and Irian Jaya).
The results are presented in Table 2.

Sex

Sex by itself had no significant effect at P<0.05 on
any individual character.

Age

Most variables showed a significant influence of
individual age; exceptions were NB, TR, IL, MSF,
BB, RAP, M'W and M J L (P<0.05).

Locality

A locality is generally an island, except that the
mainland Southeast Asian population is treated as
a single locality (Malaysia). The variables ZB, NB,
IO, IFB, BH, MW, BS, LB, and TL, were
significantly influenced by locality.

Taxonomic status of the ricefield rat, Rattus argentiventer

55

Table 2 Multiple regressions on sex, age, locality, and the interactions among these factors for Rattus argentiventer in
skull, dentary and external characters. F values are presented for the main effects and interactions (2 way and
3 way). Significance levels are *) 0.05>p>0.01, **) 0.01>p>0.001 and ***) P<0.001. For explanation of character
codes see Materials and Methods.

Dependent

Variable

Locality

Sex

Age

Locality Vs
Age

Locality Vs
Sex

Age Vs
Sex

3 way

Interactions

LB

3.530*

1.798

7.911***

2.806*

3.080*

3.190*

0.119

TL

4.494**

0.078

5.403**

1.323

0.989

1.950

0.279

E

1.486

0.298

7.077***

2.403

1.216

0.839

0.079

HF

0.836

0.022

0.791

0.070

0.267

0.281

0.143

GSL

1.681

1.846

23.601***

1.798

0.450

1.326

1.771

ZB

3.029*

1.421

16.225***

3.047*

2.278

1.802

0.008

HB

2.308

0.935

8.773***

1.606

0.930

2.938

0.121

NL

1.231

0.162

18.706***

1.082

1.114

0.932

1.184

NB

4.542**

1.574

2.476

0.393

0.707

0.012

0.963

IO

15.608***

1.045

6.579**

0.803

2.731*

0.807

0.008

BOZP

1.421

0.150

5.526**

0.422

0.569

0.410

0.058

LOP

0.126

3.213

14.552***

2.138

3.840*

0.743

0.554

TR

2.496

0.094

0.074

0.312

0.277

0.646

0.915

IL

1.082

2.642

2.316

1.022

0.198

1.512

0.389

MSF

2.679

0.021

2.914

0.658

0.659

0.320

1.251

IFB

3.252*

0.113

6.514**

0.175

2.855*

0.926

1.274

BL

2.330

0.040

8.130***

1.763

0.862

0.158

2.198

BH

3.566*

2.856

4.190*

0.383

1.332

0.676

0.630

BB

1.765

1.317

1.732

1.152

0.171

0.480

1.200

LOD

0.335

0.784

14.566***

1.942

2.991*

0.033

0.728

RAP

0.392

1.141

2.550

3.719**

0.895

1.142

0.265

DL

2.126

0.003

9.316***

1.656

0.382

0.219

1.337

M 3 W

2.804*

0.006

0.122

0.244

0.459

2.402

0.286

M’L

0.806

0.063

0.355

0.383

0.538

0.477

1.092

CBL

2.421

0.925

27.527***

2.877*

0.619

1.545

1.539

M 3 M 3

0.562

0.231

10.771***

0.980

0.446

0.272

0.346

M 2 M 2

1.380

0.224

11.644***

2.231

1.494

0.382

0.209

M 3 M 3

2.605

2.571

9.670***

2.664*

2.418

0.286

0.925

BS

3.395*

1.449

7.235***

0.715

0.108

1.018

1.628

POW

2.011

0.806

6.465**

0.851

1.569

0.731

0.036

Interaction

The variables ZB, CBL, M 3 M 3 , LB at P<0.05 and
RAP at P>0.01 were significantly influenced by the
interaction between locality and age. The characters
IO, LOP, IFB, LOD and LB (P<0.05) were
significantly influenced by the interaction between
locality and sex. Except for head and body length
(P<0.05) there was no significant influence between
age and sex (P>0.05). No variable showed a
significant interaction between all three factors.

Multivariate analysis

The regression analysis showed that age strongly
influenced many of the skull, dental and dentary
characters, either individually or through
interactions with sex or locality. Consequently the
multivariate analysis was based on fully adult
specimens as judged from basicranial fusion.

Although there is little interaction between sexual
dimorphism in R. argentiventer, a total of five characters
(LB, IO, LOP, IFB and LOD) showed a significant
difference between sex and locality. For this reason,
discriminant function (canonical variate) analysis
was performed separately for males and females.

Females

An initial DFA was run using skull, dental and
dentary characters and all locations. Because the
number of characters employed in this analysis
exceeded the sample size of the smallest group
(Sumba, N=9), the DFA was repeated using a
reduced set of seven characters in order to avoid
over fitting the data (Kitchener and Maryanto 1995).
The seven characters were selected by minimizing
Wilk's lambda on the first canonical variate. The
DFA plot was based on seven characters similar to
that based on the complete character set. The seven
characters in the DFA were IO, BH, BOZP, BS,
M 3 M 3 , NB and MSF. This analysis extracted three
significant functions, which together accounted for
100% of the variation. Discriminant Functions (DF)
1, 2, and 3 accounted for 63.02%, 24.60%, and
12.38% of the variation between populations
respectively (Table 3). Each of the functions was
highly significant DF1 (X 2 =200.411; df=21;
P=0.00001), DF 2 (X 2 =91.305; df=12; P=0.00001) and
DF 3 (X 2 =33.432; df=5; P=0.00001). Reclassification
of cases resulted in a correct allocation of 95.3% to
localities

56

I. Maryanto

DF 1

DF 1

DF 2

Figure 4 Plot of the male canonical variate on the following island group population 1; ♦ Java group (Thailand,
Malaya, Sumatra, and Java); ■ Kalimantan group (Kalimantan Island); O Bali group (Bali, Lombok,
Sumbawa, Sangeang, Komodo, Flores, Adonara, Lembata, Alor, Timor and Tanimbar) and © Sumba group
(Sumba Island). A. Discriminant Function (DF)1 vs. Discriminant Function 2. B. Discriminant Function 1 vs.
Discriminant Function 3 and C. Discriminant Function 2 vs. Discriminant Function 3.

Taxonomic status of the ricefield rat, Rattus argentiventer

Three of the seven characters (BOZP, IO and NB)
loaded heavily on DF1 (>0.5). These are evidently
major characters, which display regional
heterogeneity in populations of R. argentiventer in
Indonesia. The DFA coefficients from this analysis
are given in Table 3.

The configuration of DFA plots suggest that the
ricefield rat contains four major morphological
subpopulations (Figure 3). These are 1, a Java group
(Thailand-Malaysia-Sumatra and Java); 2,
Kalimantan; 3, a Bali-Sulawesi group (Bali, Lombok,
Sumbawa, Sangeang, Komodo, Flores, Adonara,
Lembata, Alor, Timor, Tanimbar and Sulawesi) and
4, Sumba. A bivariate plot between DF1 and DF2
shows that the Java group is separated from the
Kalimantan, Bali-Sulawesi and Sumba groups on
DF1 (Figure 3). A plot of DF2 against DF3 shows
good separation of each of the Kalimantan, Sumba
and Bali-Sulawesi groups. A single specimen from
Irian Jaya, which was unallocated in the DFA falls
within the Java group (Group 1). The Sulawesi
population falls within group 3. A total of 94.2% of
individuals in the Java group were correctly
reclassified, with 5.7% (3 individuals) misclassified
as Kalimantan. In the Bali-Sulawesi group, 94.3% of
individuals were correctly reclassified with 2
individuals (2.9%) misclassified, one to each of the
Java and Kalimantan groups. Finally, 100% (9
specimens and 10 specimens) of individuals from
each of Sumba and Kalimantan were correctly
reclassified.

Table 3 Standardized and unstandardized (in brackets)
canonical variate function coefficients the
derived from seven character analysis of
female Rattus argentiventer

Variable

Function 1

Function 2

Function 3

IO

0.888 (3.784)

-0.472 (-2.014)

0.178 (0.760)

BH

0.066 (0.252)

0.547 ( 2.087)

0.782 (2.986)

BOZP

-0.794 (-1.953)

-0.218 (-0.537)

-0.289 (-0.712)

NB

0.639 (1.991)

0.437 (1.359)

-0.503 (-1.564)

M 3 M 3

-0.306 (-0.702)

-0.781 (-1.791)

0.420 (0.963)

BS

0.389 (0.589)

0.870 (1.316)

-0.471 (-0.712)

MSF

Variation

-0.353 (-1.617)

0.133 (0.609)

0.675 (3.089)

explained

63.02%,

24.60%,

12.38%

Constant

-25.078

-20.127

-15.232

Males

A DFA for males was run initially for the full set
of 24 skull and dentary characters for all islands.
However, because the sample size of the smallest
groups (Kalimantan and Sumba) was 5 and 6, the
analysis was repeated using a reduced set of five
characters (BS, IO, NB, NL and M'M 1 ) selected as
for the female analysis.

The five characters selected by minimizing Wilk's
lambda on the first canonical variate were IO, NB,
M'M 1 , BS, and NL. Three of those variables were

57

the same as those selected in the female analysis
(IO, NL, BS); the others relate to similar components
of the skull, namely the rostrum and the palate. The
DFA plots produced with the reduced character set
are similar to those based on the complete set of
characters. This analysis extracted three significant
functions, which together explained 100% of the
variation. The DF 1: 67.93%, DF 2: 27.48%, and DF
3: 4.59% of each of these functions is highly
significant (DF1: X^=138.065; df=15; P=0.00001, DF
2: X 2 =54.464; df=8; P=0.00001 and DF 3 X^.553;
df=3; P=0.022). The characters which loaded most
heavily (>0.5) on DF1 were IO, M'M 1 and NB.

The percentage of individuals correctly
reclassified was 87.95%. In the Java group 91.5% of
individuals were correctly reclassified with only
4.3% (2 individuals) misclassified to the Sumba
group, and 4.3% to the Bali-Sulawesi group. A total
of 66.7% of the individuals from Sumba were
correctly reclassified with 33.3% (2 individuals)
misclassified to the Bali-Sulawesi group. In the Bali-
Sulawesi group, 88% of specimens were correctly
reclassified with 4% (1 individual) and 8% (2
individuals) misclassified to the Java and Sumba
groups respectively. Finally, 80% of individuals
from Kalimantan were correctly reclassified with
one individual misclassified to the Java group.

The Scatter plot between coefficient canonical 1
and 2 showed that the Java group was separated
from the Sumba and Kalimantan groups (Figure 4).
This bivariate plot also indicated that the Kalimantan
individuals were separate from those in the Java,
Bali-Sulawesi, and Sumba groups. Furthermore,
individuals from Sumba were not clearly separated
from those in the Bali-Sulawesi group.

SYSTEMATICS

Rattus argentiventer argentiventer Robinson and
Kloss, 1916

Rattus rattus braivicaudatus Horst and Raadt,
1918

Rattus rattus chaseni Sody, 1941
Holotype

British Museum (Natural History) 19.11.5.89

Type locality

Pasir Ganting, west coast of Sumatra

Diagnosis

Robinson and Kloss (1918:55) described R. a.
argentiventer as dorsally coarsely streaked, blackish
and warm buff, grayer on the sides and limbs. The
under parts are not clearly margined and are clad
with fur having gray bases and white tips
producing a general silvery effect, except on the

58

I. Maryanto

throat, which is white. A scarcely indicated huffy
median stripe is present on the chest. The feet are
parti-colored: tail brown throughout. This
description serves equally for specimens from
Thailand to Java. The skull of the typical form is
distinguished from that of the other subspecies by
the combination of a narrower interorbital region
and narrower rostrum (Figure 5c).

Distribution

Thailand, Malaysia, Sumatra, and Java, possibly
extending to Vietnam, Cambodia, Laos.

Rattus argentiv enter kalimantanensis subsp. nov.
Holotype

Museum Zoologicum Bogoriense, MZB22419,
adult male, weight 189 grams, carcass fixed in 70%
ethanol, skull separated from body, collected by
Ibnu Maryanto and M.H. Sinaga in March 2000.

Table 4 Male standard and unstandardized (in
brackets) canonical variate function
coefficients derived from the five character
analysis of male Rattus argentiventer.

Variable

Function 1

Function 2

Function 3

BS

-0.309 (-0.524)

-0.781 (-1.318)

0.458 ( 0.774)

IO

0.848 (3.782)

0.742 ( 3.308)

0.289 ( 1.292)

NB

0.675 (2.198)

-0.948 (-3.086)

-0.029 (-0.097)

NL

0.339 (0.347)

0.731 (0.750)

-0.992 (-1.018)

M'M 1

Variation

-0.680 (-2.100)

0.348 (1.075)

0.535 ( 1.650)

explained

67.93%,

27.48%,

4.59%

Constant

-20.355

0.947

-10.794

Paratypes

See appendix for list of specimens examined

Type locality

Saka Gunung, Sei Kambat, Cerbon, Barito Kuala,
South Kalimantan.

Diagnosis

Rattus argentiventer kalimantanensis females differ
from R. a. argentiventer in averaging slightly larger
in all skull dimensions except for BOZP. They differ
from individuals in the Bali-Sulawesi group (R. a.
pesticulus) in averaging slightly larger except
dimensions for TR, DL, MSF, M 2 M 2 , M 3 M 3 , POW,
IO, TL and E. They differ from individuals in the
Sumba group (R. a. saturnus) by averaging slightly
smaller except for NB, IO, BOZP, LOP, M1W, IFB,
and HF.

Rattus argentiventer kalimantanensis males differ
from R. a. argentiventer in averaging slightly smaller
except for NB, LOP, M l M\ POW, BB, M'W, BL, E,
and HF. They differ from R. a. saturnus in averaging
smaller except for RAP, M 3 W, M'M 1 , M 2 M 2 , M 3 M 3 ,

BS, POW, LOP, GSL, LB, E, and HF. They differ
from R. a. pesticulus in averaging slightly smaller
except for NB, IO, M 3 M 3 , BS, BB, PL, M’W, MSF,
BL, BH, and HF.

Scatter plots of skull characters show that the
interorbital breadth of male R. a. kalimantanmsis is
generally narrower relative to braincase breadth
than R. a. pesticulus (Figure 5a). The nasal breadth
females of R. a. kalimantanensis is generally larger
relative to zigomatic breadth of R. a. argentiventer
(Figure 5b)

Etymology

Named after the location on which the specimens
were collected.

Pelage

Ventral fur with yellowish white tipping and base
pastel gray to light gray; chest yellowish gray.
Dorsum light brown on the tip and dark gray to
purplish gray on the base, with sparse mixture of
yellowish gray hairs. Forefeet grayish brown to
yellowish brown, hindfeet grayish yellow to light
blond.

Distribution

Widely distributed in Kalimantan (see Appendix
for localities).

Rattus argentiventer saturnus Sody, 1941
Holotype

RMNH 9808 skin and skull collected by G. Stein

Type locality

Melolo, Sumba.

Diagnosis

Female R. a. saturnus from Sumba Island differ
from R. a. pesticulus from Bali, Lombok, Sumbawa,
Sangeang, Rinca, Komodo, Flores, Adonara, Timor,
Alor, Tanimbar and Sulawesi in averaging slightly
larger in nearly all skull characters except M 2 M 2 ,
M 3 M 3 , IO, and IFB. Males differ from R. a. pesticulus
in averaging slightly larger except for GSL, NL, NB,
MSF, IFB, BL, BH, BB, PL, M 3 W, CBL, BS, MSF, IFB,
BL, and BH.

Males differ from R. a. argentiventer, in averaging
slightly larger in all skull characters except LOD,
RAP, M 1 M 1 , M 2 M 2 , M 3 M 3 , ZB, HB, IO, and BOZP.
Females differ from R. a. argentiventer by averaging
slightly larger except for IFB.

Sody (1941) in describing saturnus, distinguished
it only from tali, the taxon that applies to R.
argentiventer from the Island of Bali. He noted that
the differences between the two taxa were the
length of tail relative to length of head and body
and length of toothrows.

Taxonomic status of the ricefield rat, Rattus argentiventer

59

Figure 5 Bivariate plots of the female specimens. A. interorbital breadth against braincase breadth; B. Bulla height
against breadth of zygomatic and C. interorbital breadth against nasal breadth. Groups of locality codes as
for Figure 3.

60

I. Maryanto

5 5.5 6 6.5 7

Interorbital breadth

Figure 6 Bivariate plots of the male specimens. A. interorbital breadth against nasal breadth B. interorbital breadth
against zygomatic breadth; C. Nasal breadth against zydomatic breadth. Locality group codes as for Figure 3.

Taxonomic status of the ricefield rat, Rattus argentiventer
Description

Musser (1972) described R.a. saturnus as having
coarsely salt and peppered yellowish brown upper
parts and silvery gray under parts as contrasted to
the solid brown upper parts and creamy or buffy
brown underparts of R. rattus. Juvenile and young
adults have yellowish orange ear tufts that contrast
conspicuously with the color of the head.

Distribution

Sumba Island

Rattus argentiventer pesticulus Thomas, 1921
Rattus rattus bali Kloss, 1921
Rattus argentiventer bali Kloss, 1921
Holotype

Adult female B.M. skin and skull in BMNH no.
21.2.9.11

Type locality

Laboehan Anak and Klungkung, Bali (Cotypes)

Diagnosis

Female R. a. pesticulus differ from female R. a.
argentiventer by averaging slightly larger except for
BOZP, IL, LB, and HF. The males differ by
averaging slightly larger except for BOZP, MSF, BL,
BH, PL, M 2 M 2 , and M 3 M 3 .

Description

Thomas (1921) described R. a. pesticulus from the
Sulawesi type specimen as fur thin and coarse, not
definitely spinous. General color above dull reddish
brown, sides rather grayer, under surface sharply
defined white, the hairs on the throat with gray
bases. Hand and feet white. Tail of medium length
thinly haired, light brown, almost white basally.
The color variation with Bali specimens that are
described by Kloss (1921) as dorsally grizzled
ochraceous tawny and brownish black, below
creamy white often with traces of a median grey
stripe. Forefeet brown, hind feet white, broadly
brown mesially. Skulls robust with rather short
broad rostrum, broad palatal foramina and large
bullae.

Distribution

Bali, Lombok, Sumbawa, Sangeang, Komodo,
Flores, Adonara, Lembata, Alor, Timor, Tanimbar
and Sulawesi

DISCUSSION

Although Chasen and Kloss (1928) believed
specimens of the ricefield rat from Kalimantan to be

61

identical to typical R. argentiventer from Sumatra,
the present analysis has shown that these
populations are quite distinct on skull morphology.
Differentiation on external characters is less
pronounced with poor discrimination between
these populations based on external measurements;
typically less than 50% of individuals could be
correctly classified. One external feature that does
distinguish the population from the others is the
more yellowish ventral surface due to an increased
number of ventral hairs with yellow tips in
Kalimantan specimens. This taxon is undescribed.
At present it is best referred to as the "Kalimantan
Population" of R. argentiventer. Musser (1973)
argued that individuals from Sumba closely
resemble those from Bali and allied these
populations to those found further east on Lombok,
Sumbawa, Komodo, Rinca, Flores and Timor. In the
present analysis female specimens from Sumba are
distinct from all other populations. In contrast,
males from Sumba show overlap with other groups.
I consider that R. argentiventer from Sumba are
sufficiently distinct to warrant subspecies status as
R. a. saturnus Sody, 1941.

Specimens from Sulawesi appear from this
analysis to be referable to Bali, Lombok to Timor
and the Tanimbar Islands group. Musser (1973)
considered R. argentiventer to be recently
introduced to Sulawesi and this is supported by
my findings. Thomas (1921) described specimens
from Sulawesi collected 1908 by Dr. Mohari from
Manado as Rattus pesticulus In the same year he
redescribed Rattus pesticulus. Musser (1973; 1977)
considered R. pesticulus to be a junior synonym of
R. argentiventer. Also in 1921 Kloss described R. r.
bali. Laurie and Hill (1954) considered that R. r.
bali was a synonym of R. argentiventer as did
Musser (1973). The specimens included in this
study from Lombok were referred to R.
argentiventer bali by Kitchener et al. (1990). Since
my analysis indicates that the Sulawesi population
is indistinguishable from those from Bali, Lombok,
Sumbawa-Timor and Tanimbar Islands then two
names become available for this taxon: R. bali and
R. pesticulus. Thomas described Rattus pesticulus
twice, once in May 1921 (Thomas 1921a) and once
in an undated volume of Treubia in 1921 (Thomas
1921b). In the same volume of Treubia, but with a
later pagination, Kloss described R. bali. The earlier
page number for R. pesticulus gives it
nomenclatural priority over R. bali, thus fixing the
subspecific name for the R. argentiventer rats of
Bali, Lombok, Sumba, Timor and the Tanimbar
Islands as R. a. pesticulus (Thomas, 1921).

Specimens from Java, Sumatra and Thailand
could not be separated in this study. This supports
the view that R. a. brevcaudatus (Horst & de Raadt)
from Java and R. a. chaseni (Sody) from Malaysia
are identical with typical R. argentiventer.

62

I. Maryanto

The single specimen from Tanah Merah, Irian
Jaya fitted within the scatter of R. a. argentiventer
rather than with the geographically closer R. a.
pesticulus. This is quite possible since the port at
Tanah Merah has been used by navigators and
trading vessels for centuries as a safe anchorage
(Flannery 1990).

A major limitation to examining the morphological
differences between populations of R. argentiventer is
the marked variation related to individual age. In the
present study it was necessary to separate juveniles
and subadults from mature animals and to exclude
the younger individuals from the analysis.
Furthermore it was necessary to analyse males
separately from females. For these reasons, it has
proved impossible to provide concise diagnoses for
the various morphologically distinct subspecies of R.
argentiventer I have recognised on morphometric
grounds. Kitchener and Suyanto (1996) discussed a
similar pattern of morphological variation for a
range of mammal groups in Indonesia. They
concluded that the geomorphological and climatic
diversity within the Indonesian Archipelago has
provided an ideal setting for complex evolutionary
processes leading to morphological change. They
further suggested that many of these morphological
changes might be of relatively recent origin. This
observation is particularly pertinent in the case of
taxa such as R. argentiventer, which may have
expanded its geographic range in parallel with the
spread of rice agriculture over the last few thousand
years. An analysis of genetic markers will be needed
to fully elucidate some of these recent evolutionary
events.

ACKNOWLEDGEMENTS

I am grateful to Mr. M.H. Sinaga, technician of
Balitbang Zoologi-LIPI, Dr. R.A. How and Mr. Ron
Johnstone from the Western Australian Museum,
Dr. Kelvin Lim and C.M. Yang from the University
of Singapore for assisting in the laboratory and
fieldwork. I am also grateful to Dr. D.J. Kitchener
from the Western Australian Museum, for funding
fieldwork to the Lesser Sunda Islands, for his
companionship in the field and for his comments
on this manuscript. I am grateful also to Dr. Chris
Watts from the South Australian Museum, for his
comments on the manuscript. Finally I am grateful
to the Nagao Foundation for supporting
development of data from the Zoological Reference
Collection, National University of Singapore and for
supporting fieldwork in Sumatra and Kalimantan.

REFERENCES

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Horst M.D. and de Raadt, O.L.E. (1918). De juiste namen
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Taxonomic status of the ricefield rat, Rattus argentiventer

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Robinson, H.C. and Kloss, C.B. (1918). Mammals of
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Manuscript received 17 December 2001; accepted 11 November
2002

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I. Maryanto

Appendix 1 List of locality, sex, number of specimens, latitude and longitude (ZRC = Zoological Reference Collection,
Singapore; MZB = Museum Zoological Bogoriense, Bogor-Indonesia; WAM = Western Australian
Museum, Perth Western Australia).

Malaysia and Thailand

Kualalumpur, Female, ZRC 2143, (3.08 N, 101.75 E)
Pahang, Female, ZRC 87140, 87341 (3.37 N, 102.83 E)
Perak, Female, ZRC 269/3, ZRC 3716, ZRC 4269, ZRC
4273, ZRC 6197 (4.87 N, 100.75 E)

Perak, Male, ZRC 580/1, ZRC 584/1, ZRC 909/2, ZRC
2044, ZRC 4268 (4.87 N, 100.75 E)

Bang-Penin-Siam, Female, ZRC 4274, ZRC 19073 (6.83 N,
101.25 E)

P.Sirih-Siam, Male, ZRC 4227.

Sumatra

Jabung Lampung, Male, MZB11480 (5.48 S, 105.67 E)

Kayu Agung, Female, MZB 12795 (3.40 S, 104.83 E)

Kp. Halaban, Painan, Male, MZB 4864, MZB 4870, MZB
4873 (1.35 S, 100.57 E)

Kp. Halaban, Painan, Female, MZB 4871 (1.35 S, 100.57 E)
Loboek Sikaping, Female, MZB 4862, MZB 4865, (0.13 N,
100.17 E)

Loeboek Sikaping, Male, MZB 4869, MZB 15345 (0.13 N,
100.17 E)

Memudjan, Padang, Female, MZB 4866, MZB 4868, (0.95
S, 100.35 E)

Padang, Male, MZB 15341, MZB 15347, MZB 15349 (0.95
S, 100.35 E).

Padang, Female, MZB 4872 (0.95 S, 100.35 E)

Mulyorejo, Way Abung, Female, MZB 13336, (4.83 S,
104.88 E)

Mulyorejo, Way Abung, Male, MZB 13431 (4.83 S,
104.88 E)

Pematangsiantar, Sumatra, Male, MZB 10459, MZB 10460
(2.95 N, 99.05 E)

Sindang Datar Lebuay, Pulau Panggung, S. Lampung
Female, MZB 11245 (2.80 S, 105.95 E)

Wai Lima Lampung Male, MZB 337, MZB 361, (9.48 S,
104.52 E)

Java

Banyuwangi, Male, MZB 15348, (8.00 S, 114.08 E)

Bagusan W.Java, Female, MZB 3379, MZB 3381 (6.32 S,
106.83 E)

Bagusan W.Java, Male, MZB 3380, MZB 3382, MZB 3383,
(6.32 S, 106.83 E)

Bajulmati, Male, ZRC 8680 (7.93 S, 114.42 E)

Bandung, Male, MZB 10211, (6.90 S, 107.60 E)

Bogor Male, MZB 4880, MZB 4881, MZB 4886, MZB 4889,
MZB 4898, MZB 7027, MZB 7028 (6.58 S, 106.78 E)
Bogor Female, MZB 4887, MZB 4890, (6.58 S, 106.78 E)
Boyolali, Female, MZB 10205, MZB 10207, MZB 10208
(7.53 S, 110.33 E)

Boyolali, Male, MZB 10206, (7.53 S, 110.33 E)

Cawang Jakarta, Male, MZB 3342, MZB 3392 (6.25 S,
106.87 E)

Cawang Jakarta, Female, MZB 3439, MZB 8047, MZB
8054, MZB 8055 (6.25 S, 106.87 E)

Jakarta, Female, MZB 180, (6.10 S, 106.88 E)

Tj. Priok Jakarta, Male, MZB 3281 (6.10 S, 106.88 E)
Cirebon, Female, MZB 2145, MZB 2146 (6.73 S, 108.57 E)
Gandrungmangu, Cilacap, Male, MZB 13434 (7.53 S,
108.85 E)

Garut W.Java, Female, MZB 4897, (7.22 S, 107.90 E)

Kresek penameng W.Java, Female, MZB 12541 (6.10 S,
106.65 E)

Kresek penameng W.Java, Male, MZB 12542, (6.10 S,
106.65 E)

Malang Female, MZB 4885 (7.98 S, 112.62 E)

Palimanan Cirebon, Male, MZB 587 (6.73 S, 108.57 E)
Palimanan Cirebon, Female, MZB 588 (6.73 S, 108.57 E)
Pamanukan, Subang. Female, MZB 12361, MZB 12363,
MZB 12367, MZB 12369, MZB 12561 (6.27 S, 107.82 E)
Pamanukan, Subang. Male, MZB 12364, MZB 12365 (6.27
S, 107.82 E)

Pandegglang, Female, MZB 4900, (6.30 S, 106.10 E)
Randudongkal, Pemalang, Female, MZB 12375, MZB
12376, MZB 12564, MZB 12365 (7.10 S, 109.32 E)
Randudongkal, Pemalang, Male, MZB 12377, MZB 12380,
MZB 12383 MZB 12388, MZB 12389, MZB 12398, MZB
12563 (7.10 S, 109.32 E)

Serang, N. Banten, Male, MZB 4892, MZB 4893, MZB
4894, (6.08 S, 106.08 E)

Noesa Kambangan, Male, MZB 4903, MZB 5873 (7.75 S,
108.92 E)

Noesa Kambangan, Female, MZB 4904, MZB 4905, MZB
4908, MZB 4909, MZB 8444 (7.75 S, 108.92 E).

Kalimantan

Barambai S. Kalimantan, Female, MZB 11114 (3.08 S,
114.64 E)

Barambai S. Kalimantan, Male, MZB 11115 (3.08 S,
114.64 E)

S. Kambat,Cerbon-S. Kalimantan, Male, MZB 22418, MZB
22419, MZB 22424, MZB 22429 (3.06 S, 114.70 E)

S. Kambat,Cerbon-S. Kalimantan, Female, MZB 22420,
MZB 22421, MZB 22422, MZB 22423, MZB 22425,
MZB 22432, MZB 22433, MZB 22434 (3.06 S, 114.70 E)
Singkawang, Female, MZB 4162 (0.90 N, 109.00 E)

Tepian Batang (Pasir) Samarinda, Female, MZB 6390,
MZB 6391 (0.50 S, 117.15 E).

Sulawesi

Loewoe (Kalonisatie Bone-Bone), Male, MZB 6412, MZB
6413 (2.50 S, 120.75 E)

Makasar, Female, WAM 33095, (6.28 S, 119.87 E)

Makasar, Male, WAM 33354, (6.28 S, 119.87 E)

Makasar, Female, MZB 4876, MZB 4877, MZB 4879 (5.12
S, 119.40 E)

Makasar, Sulawesi, Male, MZB 4878, MZB 2355 (5.12 S,
119.40 E)

Bali

Bali, Male, MZB 17229, MZB 17232, MZB 17236 (8.50 S,
115.27 E)

Bali, Female, MZB 17231, MZB 17233, MZB 17234, (8.50 S,
115.27 E)

Bali, Male, MZB 17237 (8.15 S, 114.43 E)

Klungkung Male, MZB 197, MZB 201 (8.53 S, 115.40 E).

Lombok

Kuta Lombok, Female, MZB 17200 (8.92 S, 116.25 E)

Swda lombok, Male, MZB 4914 (8.75 S, 116.50 E)

Sumbawa

Desa Dahu, Male, MZB 17185, MZB 17186 (8.75 S,
118.43 E)

Taxonomic status of the ricefield rat, Rattus argentiventer

65

Desa Dahu, Female, MZB 17187, MZB 17188, MZB 17191
(8.75 S, 118.43 E)

Desa teluk Santong, Female, MZB 17182, (8.73 S,
117.89 E)

Dompoe, Soembawa., Female, MZB 4916, MZB 4917,
MZB 4919, MZB 4921 (8.53 S, 118.47 E)

Sangeang

Desa Santong, Sangeang Is., Female, MZB 17184 (8.22 S,
119.01 E)

Desa Santong, Sangeang Is, Male, MZB 17183 (8.22 S,
119.01 E)

Dompoe, Soembawa, Male, MZB 4918, MZB 4920 (8.53 S,
118.47 E).

Sumba

Kananggar E. Sumba, Female, MZB 839 (10.05 S,
120.37 E)

Kambaniru, Female, MZB 4924, MZB 4939, MZB 4941,
MZB 4942 (9.65 S, 120.32 E)

Melolo, Female, MZB 4931, (9.88 S, 120.67 E)

Sumba, Male, MZB 17165, MZB 17167, MZB 17168, MZB
17175 (9.63 S, 119.53 E)

Waikabubak, Male, MZB 17172, MZB 17177, (9.63 S,
119.53 E)

Waikabubak, Sumba, Female, MZB 17176, MZB 17178,
MZB 17179 (9.63 S, 119.53 E)

Flores

Horowura Boru, Male, MZB 17195, (8.55 S, 123.65 E)
Kelimutu Woloaru, Female, MZB 17192, (8.70 S, 121.90 E)
Manggarai, Male, MZB 4923 (8.50 S, 120.25 E)

Maumere, Female, MZB 10492 (8.62 S, 122.23 E)

Mboera, Female, MZB 2396, MZB 2398 (8.57 S, 119.87 E)
Mboera, Male, MZB 2397, MZB 2399 (8.57 S, 119.87 E)

Wai Sano, Female, MZB 2401 (8.66 S, 120.51 E)

Wai Sano, Male, MZB 2402, MZB 2403 (8.66 S, 120.51 E)

Lembata

Belang Watokobu, Female, MZB 17193 (8.43 S, 123.37 E)

Adonara

Waiwerang, Horowura, MZB 17194 (8.33 S, 123.15 E)

Rinca

Rinca, Female, MZB 17196 (8.65 S, 119.67 E)

Tanimbar Is

Tanimbar, Male, WAM 43645, WAM 43653, WAM 43674,
WAM 44390 (7.61 S, 131.46 E)

Tanimbar, Female, WAM 44317, WAM 43652 (7.61 S,
131.46 E)

Timor

Camplong Male, MZB 22413, MZB 22415, (10.03 S,
123.92 E)

Camplong Female, MZB 22414, MZB 22416 (10.03 S,
123.92 E)

Gunung Mutis, Timor, Male, MZB 17201 (9.55 S,
124.22 E)

Timor, Female, MZB 16747 (10.07 S, 123.88 E)

Alor

Alor, Male, MZB 17206, MZB 17207, MZB 17210, MZB
17214, MZB 17215 (8.25 S, 124.72 E)

Alor, Female, MZB 17208, MZB 17211, MZB 17212, MZB
17213, MZB 17219, MZB 17225, MZB 17226 (8.25 S,
124.72 E)

Komodo

Female, MZB 9001 (8.60 S, 119.50 E)

Male, MZB 9002 (8.60 S, 119.50 E)

Papua

Tanah Merah, Female, MZB 4875 (2.40 S, 140.35 E)

Records of the Western Australian Museum 22: 67-74 (2003).

Second species of Mangkurtu (Spelaeogriphacea)
from north-western Australia

Gary C. B. Poore 1 and W. F. Humphreys 2

1 Museum Victoria, PO Box 666E, Victoria 3001, Australia
email: gpoore@museum.vic.gov.au

2 Western Australian Museum, Francis Street, Perth, Western Australia 6000, Australia
email: humphw@museum.wa.gov.au

Abstract - A new species, Mangkurtu kutjarra, is described, the second in this
genus from Australia, and fourth living in the Order Spelaeogriphacea in the
world. It occurs in a calcrete aquifer very close to that in which its congener
lives in the Pilbara, Western Australia. Spelaeogriphacea, like
Thermosbaenacea are thought to be relicts of a more widespread shallow-
water marine Tethyan fauna stranded in interstitial or ground-water
environments during periods of marine regression.

INTRODUCTION

Over the last few years an extraordinary diversity
of stygal animals has been recorded from arid parts
of Western Australia occupying both anchialine
ecosystems and continental groundwaters
(Humphreys, 1999, 2001), both fresh and saline. As
is the case globally, these species largely comprise
crustaceans, but a diversity of dytiscid beetles
(Cooper et al, 2002; Watts and Humphreys, 1999)
and hydrobiid gastropods is also notable.

Amongst this fauna, Mangkurtu mityula Poore and
Humphreys, 1998, from the Millstream (Western
Fortescue Plain) aquifer in the Pilbara, was the first
record of Spelaeogriphacea from Australia. This
order was first described ( Spelaeogriphus lepidops
Gordon, 1957) from a sandstone cave at an altitude
of 700 m on Table Mountain, South Africa. This
remained the only record until another member of
the order was found in a cave in the Mato Grosso in
western Brazil ( Potiicoara brasiliensis Pires, 1987).
The Millstream species is the only species known to
be widely present in a major aquifer where it is
found together with a range of other Gondwanan
freshwater lineages (Poore and Humphreys, 1998).
The occurrence of fossil species of purported
spelaeogriphaceans, Acadiocaris novascotica
(Copeland, 1957) in Carboniferous marine
sediments in Canada (Schram, 1974), and in
lacustrine deposits of Jurassic age in China
(. Lianongogriphus quadripartitus Shen et al, 1998)
suggests that invasion of fresh water occurred prior
to the dissolution of Pangaea and long before the
fragmentation of Gondwana (Poore and
Humphreys, 1998).

A Tethyan fauna, allied to the stygofauna of the
Cape Range peninsula and Barrow Island

(Humphreys, 2000), intrudes towards Millstream
from the coast and is present in aquifers of the
lower reaches of the Robe and Fortescue Rivers.
Elements of this fauna reach an altitude of c. 300 m
in the Robe River (Humphreys, 2001), the
approximate level of the Late Eocene sea-level high
in south-western Australia (G.W. Kendrick,
personal communication 1999).

Here, a second species of the Australian genus is
described from material collected in the same
region as the first. Its environment is described.
Sampling was conducted and water quality
measured as detailed elsewhere (Watts and
Humphreys, 2000). Material is lodged in the
Western Australian Museum, Perth (WAM) and
Museum Victoria, Melbourne (NMV).

Order Spelaeogriphacea Gordon, 1957
Family Spelaeogriphidae Gordon, 1957
Mangkurtu Poore and Humphreys, 1998
Remarks

Mangkurtu is unique among spelaeogriphacean
genera in the possession of a digitiform
maxillipedal epipod and in having endopodal lobes
on the pleopods. The new species is similar in these
respects and in many others to M. mityula, type
species, but differs in the proportions and shapes of
some morphological features. We do not reillustrate
mouthparts but microscopic examination and SEMs
provided to us by G.D.F. Wilson show no important
differences. Mangkurtu is the only spelaeogriphid
genus with more than one species.

In Poore and Humphreys' (1998) diagnoses of the

68

G.C.B. Poore, W.F. Humphreys

three extant genera the length of the antennal 2
scale was compared to peduncular article 3. It is
now apparent that this "article 3" is in fact articles 3
and 4 together. No authors, including ourselves,
have illustrated the very short article 3 of
spelaeogriphaceans and have treated the antenna 2
peduncle as having four articles. We were able to
confirm a similar very short article 3 visible only
ventrally in a specimen of Spelaeogriphus.
Spelaeogriphaceans have five antennal articles like
all eumalacostrans (Figure lc). We are now able to
confirm that oostegites occur on pereopods 1-5 on
this species as in other spelaeogriphaceans.
Juveniles of the new species possess a reduced
pereopod 7 (Figure le) and we assume that this
manca stage, hitherto unreported, is general for
spelaeogriphaceans. Embryos flex dorsally.

Gordon's (1957) figures of maxilla 2 of
Spelaeogriphus lepidops suggest a pattern of setation
on the outer endites different from that in
Mangkurtu. We were able to observe that the
triangular setae drawn by her are in fact just the
tips of much longer curved setae, similar to those
seen in the other genera.

Mangkurtu kutjarra sp. nov.

Figures 1-3

Material examined

Holotype

Western Australia. Pilbara Region, Fortescue
River Valley, Roy Hill Station, Battle Hill Well,
22°44’28"S, 120°07’37"E, W.F. Humphreys and J.M.
Waldock, 8 September 2000 (stn BES 8505), WAM
C29101 (male, 4. 5. mm, with 5 slides).

Paratypes

Collected with holotype, WAM C29102 (female
with oostegites, 4.5 mm) C29103 (female with
oostegite buds, 3. 6. mm, with 3 slides), C29104
(juvenile, 4.0.mm), C2915 ( 2 males, 4.0, 4.6 mm),
C29106 (female with oostegites and 16 eggs),
C29107 (4 females, incomplete), C29108 (20
juveniles and mancas, incomplete); NMV J57461
(male), J57462 (male, 4.5 mm), J57463 (female with
oostegite buds, 3.6 mm), J57464 (male, 4.2 mm),
J57465 (male, 4.2 mm), J57466 (juvenile). Same area
and date. Aerodrome Bore, 22°42'53"S, 119°54'53"E
(stn BES 8501), NMV J57467 (male, 2 females, 2
juveniles).

Description

Male. Ratio of carapace : pereon : pleon ratio 1 :
1.9 : 3.2. Telson 1.2 times as long as wide at base,
tapering to semicircular apex with three pairs of
marginal spiniform setae, first pair (most lateral)
the longest; and 1 shorter pair submarginal-dorsal

between 2 most lateral setae.

Antenna 1 total length 0.65 of body; outer
flagellum with c. 37 articles; inner flagellum of 30
articles.

Antenna 2 1.5 times length of body; peduncle
article 2 with distolateral row of stout setae; article
3 very short; article 4 longer than article 2; article 5
longer than 4, with distal setae; exopod width 0.30
length, with medial rows of alternating eight
plumose and eight stout setae.

Pereopods 1-7 similar to those of M. mityula,
endopod and exopods of similar proportions, bases
of 5-7 with similar pedunculate setae. Distal
margins of carpi of all pereopods with oblique row
of falcate setae mounted on conical projections: nine
or ten on pereopods 1-3, four-seven on pereopod 4;
propodi with groups of one-three similar setae.

Penes at bases of pereopods 7 cylindrical, stout,
meeting in midstemum.

Pleopod 2 exopod two-articulate, shorter than
endopod; proximal article with one lateral simple
seta, one distolateral plumose and one distomesial
plumose setae; distal article with 18 marginal
plumose setae and 13 shorter simple setae on distal
margin; endopod broadly-falcate, curving laterally
to enclose exopod, with one proximomesial and 4
midmesial setae.

Uropod peduncle with four distolateral, 2
distomesial, one proximomesial spiniform setae,
one simple and one plumose setae ventrally on
distal margin; endopod with 22 marginal plumose
setae and two or three submarginal dorsal setae;
exopod two-articulate; proximal article longer than
endopod, with four distolateral, six mesial
spiniform setae; distal article twice as long as wide,
with 26 marginal plumose setae, one simple apical
setae.

Manca. Pereonite 7 apparent as a single article.

Ovigerous female. Ratio of carapace : pereon : pleon
ratio 1 : 2.5 : 2.7. Pleopod 2 similar to pleopods 1
and 3; oostegites on pereopods 1-5, carrying in one
case 16 eggs and in another four embryos. Embryos
flex dorsally.

Etymology

Kutjarra, meaning two in the Australian
Aboriginal language of the Western Desert,
alluding to the second species in the genus.

Distribution

Roy Hill calcrete, Pilbara, Western Australia.
Remarks

Mangkurtu kutjarra differs from M. mityula in the
shape of the telson (more tapered and with a more
rounded apex, four rather that five pairs of distal
marginal setae), proportions of the uropod
(endopod shorter and relatively wider, with more

Second species of Mangkurtu

69

Figure 1 Mangkurtu kutjarra n. sp. Holotye male, 4.5 mm, WAM C29101. a, b, habitus (scale 0.5 mm)- c antenna 2
peduncle; d, telson. Manca, WAM C29108. e, (right to left) pereonites 6 and 7, pleonite 1 with basis of
pereopod 6, pereonite 7, and peduncle of pleopod 1.

70

G.C.B. Poore, W.F. Humphreys

Figure 2 Mangkurtu kutjarra n. sp. Holotype male, 4.5 mm, WAM C29101. a-d, pereopods 1, 2, 4 and 6 with details of
distal articles; e, ventral view of pereonite 7 with penes.

Second species of Mangkurtu

71

Figure 3

Mangkurtu kutjarra n. sp. Holotype male, 4.5 mm, WAM C29101. a, uropod; b, pleopod 2.

setae), and shape of the male pleopod 2 endopod
(much more curved and broader).

The environment

The Fortescue Valley extends for 450 km as a
broad plain trending west-north-west between the
Hamersley and Chichester Ranges in the Pilbara,
the northern craton of the 'Western Shield' of
Australia. The upper part of the river and its
tributaries drain the Archaean ranges to the
Cainozoic surficial alluvial deposits in the Fortescue
Valley.

The Fortescue Valley contains several discrete
groundwater calcrete deposits, the larger being
those at Roy Hill and the Western Fortescue Plain
(Millstream, a lacustrine calcrete of Tertiary age) —
the nature of these calcretes is discussed elsewhere
in the context of stygofauna (Humphreys, 1999;
2001). Further calcrete bodies occur downstream of
Roy Hill, successively at Mulga Downs, Mount
Florance and Millstream, the latter being the habitat
of Mangkurtu mityula, before the river descends
through groundwater gaps to the coastal plain.

Sampling has not detected spelaeogriphaceans in
the Mulga Downs and Mount Florance calcretes,
although other stygofauna do occur there.

Mangkurtu kutjarra occurs in a groundwater
calcrete aquifer on Roy Hill pastoral station in the
upper Fortescue Valley (Figure 4), immediately
upstream of the Fortescue Marsh which generally
comprises a salt lake (playa). The Fortescue Marsh
is now an internal drainage basin within the
Fortescue palaeodrainage and even in times of peak
flow - following rare episodic rainfalls, when an
extensive lake may form lasting many months to
several years - it does not discharge downstream.
Consequently, the area accumulates salt from aerial
deposition and contains hypersaline groundwater.
Floods that do recharge the Millstream aquifer
downstream result from flows of the South
Fortescue River. Hence, the Millstream aquifer is
now probably isolated from those upstream in the
Fortescue Valley, including the Roy Hill calcrete. In
the Eocene the Fortescue Marsh drained externally
through the current Robe River and a variety of
evidence, though not conclusive, suggests that the

72

G.C.B. Poore, W.F. Humphreys

Figure 4 Schematic diagram of the Fortescue Valley and the spatial relationship of the Roy Hill calcrete area to other
calcretes in the drainage system and the Fortescue Marsh.

Marsh could have been essentially internally
draining since the Miocene. However, the
groundwater system is still continuous and the
amount of salt in the hypersaline groundwater in
the Marsh suggests that there have been exceptional
wet periods in which the marsh overflowed to the
west (D.P. Commander, personal communication
2003).

The water inhabited by Mangkurtu kujarra is
slightly saline and with low oxygen saturation but
with a pH typical of calcrete areas (Table 1). It is not
known whether the water column is highly
stratified with respect to salinity as found in some
other arid areas of Western Australia (Watts and
Humphreys, 2000; Humphreys, 2002).

BIOGEOGRAPHIC AND CONSERVATION
IMPLICATIONS

The presence of isolated populations of separate
species of spelaeogriphaceans suggests that the
groundwater calcrete deposits have long been
separated from each other. Similar speciation is
seen amongst stygal Dytiscidae (Insecta:
Coleoptera) in the Yilgam (the southern part of the
Western Shield) where both morphological and
molecular evidence support the thesis that the
calcretes form separate islands of groundwater.

Table 1 Site description and water quality at two
collection sites from which M. kutjarra was
taken at Roy Hill Station, Fortescue River
Valley, Pilbara Region, WA.

Parameter

Battle Hill

Well

Aerodrome

Bore

Site description

Well maintained,
uncovered, cement-
lined well on raised
stone plinth

Overgrown,
open borehole,
fetid

Conductivity (mS cm ') 6.65

5.67

Salinity (g L ’)

3.94

3.36

Temperature (°C)

25.5

29.2

0 2 (mg L>)

1.5

0.71

O saturation (%)

18.6

9.2

pH

7.13

7.20

Depth to water (m)

3.6

9

Depth of water (m)

4.0

2.0

many containing a unique fauna (Watts and
Humphreys, 1999, 2000, 2001; Cooper et a]., 2002).
This appears to be the case there also with stygal
Haloniscus (Crustacea: Isopoda: Oniscidea) which
occurs as sympatric congeners (Taiti and
Humphreys, 2001).

The Fortescue calcrete aquifers contain numerous
new genera and species of Candoninae (Crustacea:

Second species of Mangkurtu

73

Ostracoda): one an undescribed genus from the Roy
Hill calcrete as yet unknown elsewhere, others
species from the calcretes upstream (Ethyl Gorge)
representing two different undescribed genera,
while a further two undescribed genera occur
downstream of Roy Hill, from Mulga Downs to the
coast. Furthermore, a tributary joining the central
Fortescue Marsh (Weeli Wolli Creek) contains a
further two undescribed genera (I. Karanovic,
personal communication). In addition, isopods of
the genus Pygolabis, second genus of the enigmatic
family Tainisopidae first described from the
Kimberley (Wilson and Ponder, 1992), are
represented by at least four species (Wilson, 2003)
in separate calcretes found in drainages to the main
Fortescue Valley.

The occurrence of distinct faunas in the discrete
deposits of groundwater calcrete on the Western
Shield has become contentious owing to the
conservation implications of numerous discrete
faunas in shallow unconfined groundwaters in an
arid but mineraliferous region. For example,
numerous species of Chydaekata (Amphipoda:
Paramelitidae) have been described from the
calcretes of the upper Fortescue River, at Ethyl
Gorge (Bradbury, 2000) but thus far these have not
been corroborated by electrophoretic data. Among
the diverse copepod fauna of the Yilgarn
(Karanovic, in press), many species have quite
widespread distributions, not being confined to
single calcretes. It is not unexpected to find a
diversity of responses amongst lineages of different
ages and ecologies in response to the evolution of
this ancient landscape.

Biogeographic implications

Present-day Thermosbaenacea (which comprise
only 34 species in total) are thought to be relicts of a
more widespread shallow-water marine Tethyan
fauna (Wagner, 1994), stranded in interstitial or
ground-water environments during periods of
marine regression. The same hypothesis has been
applied to other taxa such as stygiobiont
Amphipoda (Stock, 1993) and the copepod genus
Misophriopsis (Boxshall and Jaume, 2000). This
hypothesis can be extended to the four
spelaeogriphacean species and the existence of a
supposed marine fossil, Acadiocaris novascotica
(Copeland, 1957) from the Carboniferous in Canada
(Schram, 1974), gives some support.

This is consistent with the intrusion of Tethyan
fauna, especially Halosbaena tulki
(Thermosbaenacea), up the Robe River, the fomer
Fortescue drainage and close to Mangkurtu mityula
sites, to an altitude of c. 300 m (Humphreys, 2001),
the approximate level of the Late Eocene sea-level
high in south-western Australia (G.W. Kendrick,
personal communication 1999). However, the three
extant genera of Spelaeogriphacea all occur on parts

of the continents that have not been inundated by
the sea since the dissolution of Gondwana, which
commenced with the separation of east- and west-
Gondwana (142-133 Ma: Partridge and Maud,
1987). Each species now occurs with a very
circumscribed distribution in subterranean
freshwater habitats on Gondwanan fragments
(Africa, South America and Australia) (Poore and
Humphreys, 1998). While both Mangkurtu and
Spelaeogriphus locations may have abutted the
shoreline of Cretaceous or Late Jurassic marine
incursions (Scotese, 1998), such proximity in the
case of Potiicoara in the southern Mato Grosso,
Brazil, is obscure. The sites lie in an extensive karst
area cut by the Rio Paraguai valley at an altitude of
only c. 330 m, despite lying nearly 1000 km inland,
and the cave waters inhabited descends about 200
m (Moracchioli, 2002). Hence the area may have
abutted Eocene shores and a clear marine intrusion
into the region in the Late Miocene (10 Ma) and less
clearly in the Palaeocene (60 Ma) directly from
western Tethys was shown by Smith et al. (1994).

ACKNOWLEDGEMENTS

We thank the lessees of the pastoral stations for
facilitating access and Julianne Waldock for
wonderful support in the field and the laboratory.
For specimens of the extant non-Australian
Spelaeogriphacea we thank Nicoletta Moracchioli
and Norma Sharratt. We thank G.D.F. (Buz) Wilson,
Australian Museum, Sydney for allowing us to see
the SEMs prepared by him. Field work was funded
by the BankWest Landscape Conservation Visa Card
Trust Fund Grants.

REFERENCES

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misophioirds (Crustacea: Copepoda) shed new light
on the origin of anchialine faunas. Zoologischer
Anzeiger 239: 1-19.

Bradbury, J.H. (2000). Western Australian stygobiont
amphipods (Crustacea: Paramelitidae) from the Mt
Newman and Millstream regions. Records of the
Western Australian Museum, Supplement 60: 1-102.

Cooper, S., Hinze, S., Leys, R., Watts, C.H.S. and
Humphreys, W.F. (2002). Islands under the desert:
molecular systematics and evolutionary origins of
stygobitic water beetles (Coleoptera: Dytiscidae) from
central Western Australia. Invertebrate Systematics 16
589-598.

Copeland, M.J. (1957). The Carboniferous genera
Palaeocaris and Euproops in the Canadian maritime
provinces, journal of Paleontology 31: 595-99.

Gordon, I. (1957). On Spelaeogriphus, a new cavernicolous
crustacean from South Africa. Bulletin of the British
Museum of Natural History (Zoology) 5: 31-47.

Humphreys, W.F. (1999). Relict stygofaunas living in sea
salt, karst and calcrete habitats in arid northwestern

74

G.C.B. Poore, W.F. Humphreys

Australia contain many ancient lineages. In: Ponder,
W.F., and Lunny, D. (eds). The other 99%. The
conservation and biodiversity of invertebrates. Royal
Zoological Society of New South Wales, Mossman.
Pp. 219-227.

Humphreys, W.F. (2000). The hypogean fauna of the
Cape Range peninsula and Barrow Island,
northwestern Australia. In: Wilkins, H., Culver, D.C.,
and Humphreys, W.F. (eds). Ecosystems of the World.
Elsevier, Amsterdam. Pp. 581-601.

Humphreys, W.F. (2001). Groundwater calcrete aquifers
in the Australian arid zone: the context to an
unfolding plethora of stygal biodiversity. In:
Humphreys, W.F. and Harvey, M. S. (eds).
Subterranean Biology in Australia 2000. Records of the
Western Australian Museum, Supplement 64: 63-83.

Humphreys, W.F. (2002). Groundwater ecosystems in
Australia: an emerging understanding. Proceedings of
the International Association of Hydrogeologists
Conference, Darwin, Australia 12-17 May 2002. CD-
ROM.

Karanovic, T. (in press). Subterranean copepods
(Crustacea: Copepoda) from arid Western Australia.
Crustaceana Monographs 3.

Moracchioli, N. (2002). Estudo dos Spelaeogriphacea
brasileiros, crustaceos Peracarida subterraneos.
Doutor em Ciencias thesis. Institute de Biociencias da
Universidade de Sao Paulo, Sao Paulo.

Partridge, T.C. and Maud, R.R. (1987). Geomorphic
evolution of southern Africa since the Mesozoic. South
African Journal of Geology 90: 179-208.

Pires, A.M.S. (1987). Potiicoara brasiliensis: a new genus
and species of Spelaeogriphacea (Crustacea:
Peracarida) from Brazil with a phylogenetic analysis
of the Peracarida. Journal of Natural History 21: 225-
238.

Poore, G.C.B. and Humphreys, W.F. (1998). First record
of Spelaeogriphacea from Australasia: a new genus
and species from an aquifer in the arid Pilbara of
Western Australia. Crustaceana 71: 721-742.

Schram, F.R. (1974). Paleozoic Peracarida of North
America. Fieldiana Geology 33: 95-124.

Scotese, C.R. (1998) Paleogeographic Atlas. PALEOMAP
Project. University of Texas, Arlington.

Shen, Y.-b., Taylor, R.S. and Schram, F.R. (1998). A new
spelaeogriphacean (Crustacea: Peracarida) from the
Upper Jurassic of China. Contributions to Zoology 68:
19-35.

Smith, A.G., Smith, D.G. and Funnell, B.M. (1994). Atlas
of Mesozoic and Cenozoic coastlines. Cambridge
University Press, Cambridge.

Stock, J.H. (1993). Some remarkable distribution patterns
in stygobiont Amphipoda. Journal of Natural History
27: 807-819.

Taiti, S. and Humphreys, W.F. (2001). New aquatic
Oniscidea (Crustacea: Isopoda) from groundwater
calcretes of Western Australia. Records of the Western
Australian Museum, Supplement 64: 133-151.

Wagner, H.P. (1994). A monographic review of the
Thermosbaenacea (Crustacea: Peracarida). Zoologische
Verhandelingen, Leiden 291: 1-338.

Watts, C.H.S. and Humphreys, W.F. (1999). Three new
genera and five new species of Dytiscidae
(Coleoptera) from underground waters in Australia.
Records of the South Australian Museum 32: 121-142.

Watts, C.H.S. and Humphreys, W.F. (2000). Six new
species of Nirridessus and Tjirtudessus (Dytiscidae;
Coleoptera) from underground waters in Australia.
Records of the South Australian Museum 33: 127-144.

Watts, C.H.S. and Humphreys, W.F. (2001). A new genus
and six new species of Dytiscidae (Coleoptera) from
underground waters in the Yilgarn palaeodrainage
system of Western Australia. Records of the South
Australian Museum 34: 99-114.

Wilson, G.D.F. (2003). A new genus of Tainisopidae fam.
nov. (Crustacea: Isopoda) from the Pilbara, Western
Australia. Zootaxa 245: 1-20.

Wilson, G.D.F. and Ponder, W.F. (1992). Extraordinary
new subterranean isopods (Peracarida: Crustacea)
from the Kimberley region. Western Australia.
Records of the Australian Museum 44: 279-298.

Manuscript received 23 April 2003; accepted 18 June 2003

Records of the Western Australian Museum 22: 75-80 (2003).

A new species of ingolfiellid amphipod (Crustacea: Amphipoda)

from Western Australia

Susana Gallego Martinez and Gary C. B. Poore

Marine Biology Section, Museum Victoria, GPO Box 666E, Melbourne, Victoria 3001, Australia
emails: sgallego@museum.vic.gov.au; gpoore@museum.vic.gov.au

Abstract - A third species, Ingolfiella quokka sp. nov., is added to the
Australian fauna, the first from Western Australia. The new species is from
an intertidal sandy beach environment; previously described Australian
species inhabit the marine shelf of south-eastern Australia.

INTRODUCTION

The ingolfiellidean amphipods are a small group
of about 30 species included in two families, the
monotypic Metaingolfiellidae, and Ingolfiellidae.
Their distribution ranges from the deep sea through
shallow marine sediments and intertidal sands to
freshwater springs and caves. The presence of an
"eye-lobe" distinguishes the suborder Ingolfiellidea
from other suborders of the order Amphipoda.
While it is probable that the group is monophyletic
their phylogenetic isolation from Gammaridea in
particular is questionable (Lowry and Poore, 1989;
Dahl, 1977). Stock (1979; 1977; 1976), Dojiri and Sieg
(1987), and Ruffo and Taglianti (1989) have studied
the systematics and zoogeography of the group.
Stock (1976) divided the family Ingolfiellidae into
three genera, and the largest genus, Ingolfiella, into
subgenera. The question of subgenera was
addressed again by Ruffo and Taglianti (1989).
Lowry and Poore (1989) had difficulty placing their
two south-eastern Australian species
unambiguously in any subgenus and questioned
their usefulness. We do not add further to this
debate, especially as the Ingolfiellidea as a whole
are subject to a phylogenetically based
reclassification (R. Vonk and F. Schram, personal
communication, 2002).

A new species of ingolfiellids was discovered in
samples taken on a sandy beach on Rottnest Island,
Western Australia, and is here described. The
material comprises 16 individuals, 2 males and 14
females. The description is a composite derived
after dissection of several individuals. Types are
lodged in the Western Australian Museum, Perth
(WAM), and Museum Victoria, Melbourne (NMV).
Abbreviations are as follows: Al, A2, antennae 1, 2;
Gl, G2, gnathopods 1, 2; MX1, MX2, maxillae 1, 2;
MD, mandible; MP, maxilliped; P3-P7, pereopods
3-7; PL1-PL3, pleopods 1-3; r, right; 1, left; UR1-
UR3, uropods 1-3.

Ingolfiellidae Hansen, 1903

Ingolfiella quokka sp. nov.

Figures 1-3

Holotype

Female, 1.08 mm (WAM C33548 on 3 slides), City
of York Bay (32°00'S, 115°36'E), Rottnest Island,
Western Australia, sandy beach, S. Griffin, July
1991.

Paratypes

Female, 1.54 mm (WAM C33549); male, 1.25 mm
(WAM C33550); female, 1.48 mm (WAM C33551,
with 2 slides); female, 1.27 mm (WAM C33552);
female, 1.08 mm (WAM C33553, with 1 slide); male,
1.06 mm (WAM C33554, with 2 slides); female, 0.87
mm (WAM C33555); female, 1.34 mm (WAM
C33556); female, 1.38 mm (WAM C33557); female,
1.31 mm (WAM C33558); female, 0.91 mm (WAM
C33559); female, 1.06 mm (WAM C33560); female,
1.26 mm (NMV J52442); female, 1.22 mm (NMV
J52443); female, 0.78 mm (WAM C33561). All
collected with holotype.

Description

Female (based on holotype, 1.08 mm, and paratype
female, 1.48 mm). Body segments laterally
compressed. Head, anterodorsal margin rounded,
without rostrum; "eye-lobe" semicircular. Pereonite
1 about half as long as head; posteroventral margin
oblique; deeper anteriorly than posteriorly such that
pereonites 1 and 2 only weakly separated.
Pereonites 2-7 increasing in depth posteriorly.
Pleonites 1-3 with posteriorly rounded epimera.
Urosomites 1 and 2 not markedly differentiated
from pleonites, of similar length; urosomite 3 with
lateral plates enclosing base of telson and uropod 3.

Antenna 1, peduncular article 1 as long as head;
ratio of articles 1.0:0.4:0.3; flagellum of four articles,

76

S. Gallego Martinez, G.C.B. Poore

Figure 1 Ingolfiella quokka sp. nov., holotype female, WAM C33548; Rottnest Island, Western Australia.

slightly more than half length of peduncle, article 4
with one apical aesthetasc; accessory flagellum of
three articles, last minute, reaching end of article 2
of flagellum. Antenna 2, peduncle as long as
peduncle of antenna 1; ratio of articles
0.4:0.2:1.0:0.8:0.8; flagellum of five articles, slightly
less than half length of peduncle, article 5 with one
apical aesthetasc.

Mandible molar processes acute; spine row of two
club-like spines; lacinia mobilis about two-thirds
width of incisor process on left, half on right, each
obscurely toothed; incisor process with five blunt
teeth on left and right. Maxilla 1, inner plate blunt,
with one seta; outer plate with four cuspidate setae,
with three, two, three, five cusps respectively; palp
of two articles, with two unequal apical setae.
Maxilla 2, inner and outer plates each with four
terminal setae. Maxilliped basal endite with two
apical setae; palp articles 1-4 with three, one, one
and one mesial setae respectively, article 5 with
long falcate unguis, seta at midlength and at base of
unguis.

Gnathopod 1 coxa at anterior of pereonite; carpus
2.4 times as long as wide, palm with three proximal
spiniform setae, three pairs of pinnate setae at mid-

palm, palm without teeth; dactylus with four teeth.
Gnathopod 2 palm at 45° to longitudinal axis;
carpus 1.7 times as long as wide; palm defined
proximally by an obtuse angle bearing two
spiniform setae (proximal one longer and more
curved than other), three triangular teeth along
palm, three flagellate spiniform setae and five
simple setae; dactylus as long as palm, with four
teeth on inner margin.

Pereopods 3 and 4, propodus with three distal
setae; dactylus with two distal setae and cylindrical
trifid unguis. Pereopods 5 and 6, merus with
spiniform seta and simple seta; carpus with two
distal long stout setae, two short stout and two
slender setae; propodus with five distal setae on
pereopod 6, and three setae on pereopod 5; dactylus
with cylindrical bifid unguis. Pereopod 7, merus
with one distal stout seta and two slender setae;
carpus with one distal short stout seta, three long
stout setae and three slender setae; propodus with
three distal setae; dactylus stout, curved with one
subtriangular spike at midlength, unguis not
defined. Coxal gills ovate, small, on pereonites 3, 4,
and 5. No oostegites.

Pleopods 1-3 subtriangular, without distal setae.

A new ingolfiellid amphipod

77

Figure 2 Ingolfiella quokka sp. nov. Paratype female, WAM C33551: A, left gnathopod 1; B, right gnathopod 2, lateral;

C, left gnathopod 2 palm and dactylus, mesial; D, pereopod 3; E, pereopod 4; F, pereopod 5; G, pereopod 6;
H, pereopod 7. Paratype male, WAM C33554: 1, right gnathopod 2, palm and dactylus, mesial.

Uropod 1 peduncle with two distal setae,
without a long reversed seta on lower margin;
inner ramus 0.9 times length of peduncle, with
lateral row of eight long setae, edges of apex finely
denticulate; outer ramus half as long as inner
ramus, with one subdistal seta. Uropod 2 peduncle
with three obliquely transverse rows of (proximal
to distal) nine, seven, nine complex setae; rami
uniarticulate, outer 0.7 times length of peduncle,
inner ramus 0.9 times length of outer ramus; outer
ramus with three setae; inner ramus with two
setae. Uropod 3 with one ramus; peduncle with

three lateral setae, ramus short, broad, with long
distal seta. Telson subsemicircular, with pair of
long dorsal setae.

Male (based on paratype, 1.06 mm). As female
except in the following. Gnathopod 2 carpus palm
defined proximally by an obtuse angle bearing one
long curved spiniform seta and more distally on
inner surface one strong flaring tooth-like seta; palm
with three subtriangular teeth and nine simple
setae. Pleopods slightly narrower than in female;
pleopod 1 only with two short distal setae. Uropod
2 peduncle with a short curved seta hooked distally

78

S. Gallego Martinez, G.C.B. Poore

Figure 3 Ingolfiella quokka sp. nov. Paratype female, WAM C33551: A, uropod 1; B, right uropod 2, lateral; C, right
uropod 2, mesial; D, detail complex setae on mesial face of uropod 2; E, antenna 1; F, antenna 2; G, maxilla 1;
H, maxilla 2; I, maxilliped; J, mandible; K, ventral view of mandibles, anterior at top, lacinia mobilis on left
and right stippled. Paratype male, WAM C33554: L, right uropod 2; M, pleopods 1-3; N, urosomite 3, dorsal
view with telson and uropods 3.

A new ingolfiellid amphipod

79

to meet a broad triangular marginal projection of
the lower margin.

Etymology

For the Quokka ( Setonix brachyurus), a species of
wallaby living on Rottnest Island in Western
Australia and after which the island was named by
Willem de Vlamingh in 1696.

DISCUSSION

Ingolfiella quokka is most similar to species that
have been assigned to the subgenus Antilleella
sharing characters such as a reduced or absent
ocular lobe, subtriangular pleopod 1 modified in
males with two apical setae, and similar pereopods
3-7. Of the species placed in this subgenus by Ruffo
and Taglianti (1989) it is similar to I. putealis, I.
fontinalis, I. tabularis, I. margaritae, I. similis, I.
unguiculata and I. beatricis as follows: (1) possession
of small ocular lobe (reduced or vestigial in most
species of the subgenus, except for I. similis where it
is absent and I. beatricis where it is developed); (2)
antenna 1 flagellum of four articles and with three-
articled accessory flagellum; (3) serrated palmar
margin of gnathopod 2 (variously eight teeth in I.
similis, smooth margin in I. beatricis); (4) gnathopod
2 sexually dimorphic; (5) gnathopods 1 and 2
dissimilar; (6) basofacial hook on peduncle uropod
2 in males; (7) subtriangular pleopods present
(subtrapezoidal in I. beatricis and I. unguiculata),
modified in males with two distal setae; and (8)
uropod 2 with three obliquely transverse rows of
spines (except for I. putealis with four rows and I.
fontinalis with two rows).

Ingolfiella quokka differs from most species of the
subgenus Antilleella in: (1) absence of a reversed
modified seta in male gnathopod 2; (2) females
without oostegites (except for I. unguiculata and I.
beatricis) - unless supposed females of our and these
species are not mature; (3) pereopods 3 and 4 unlike
pereopods 5-7, that is, dactyli of pereopods 3 and 4
with long apically bifid unguis, dactyli of
pereopods 5 and 6 with stronger bifid unguis, and
pereopod 7 lacks unguis; (4) absence of a sharp
inner spur distally on the basal part of dactyli; and
(5) dactylus of gnathopods 1 and 2 with four
serrations instead of three (except for I. beatricis).

The new species differs from I. bassiana, the only
Australian species which could be placed in
subgenus Antilleella. The complexity of the palmar
spines on the male gnathopod 2, the plumose seta
on the peduncle of uropod 1, and the two setae of
the pleopod 2 in males sets I. bassiana apart from I.
quokka.

Ingolfiella quokka has also some characters similar
to I. fuscina Dojiri and Sieg, 1987, a species more
akin to the subgenus Tethydiella Ruffo and Taglianti,
1989. These are: (1) gnathopod 1 carpus palm

bearing Y-shaped setae; (2) gnathopod 1 dactylus
with four teeth; (3) pereopods 3-4 unlike pereopods
5-7 (diagnostic of Tethydiella); (4) pereopods 3 and 4
dactyli with trifid unguis (unlike Antilleella, where
the unguis is bifid); (5) absence of a sharp inner
spur distally on the basal part of in pereopods 3-7
dactyli; and (6) the complex nature of the mesial
spines on uropod 2. However, the new species can
be distinguished from I. fuscina by: (1) absence of
unguis in pereopod 7; (2) female probably devoid
of oostegites (always present in Tethydiella); (3)
sexual dimorphism in gnathopod 2; (4)

subtriangular pleopods present in both sexes
(lamelliform with one or two apical setae in I.
fuscina); (5) male pleopod 1 subtriangular with two
apical setae (only one in I. fuscina); (6) absence of
terminal, dorsal, pectinate spine on male uropod i
(uropod 1 sexually dimorphic in I. fuscina); and (7)
presence of a basofacial hook on the peduncle of
uropod 2 in males.

Ingolfiella quokka differs from I. australiana Lowry
and Poore, 1989, another Australian ingolfiellid.
This species shares characters with species
belonging to the subgenus Trianguliella Stock, 1976.
Both species have (1) a small semicircular eyelobe;
(2) antenna 2 flagellum of five articles; (3)
accessory flagellum of antenna 1 of three articles,
the last one minute; (4) dactylus of gnathopod 1
with four serrations; and (5) pleopods 2 and 3
subtriangular in both sexes. However, I. australiana
is quite different from I. quokka : (1) antenna 1
flagellum of five articles; (2) maxilla 2, inner and
outer plates with five plumose setae; (3)
gnathopod 2 palm with distal triangular tooth and
quadrate tooth separated by a notch; (4) pereopods
5 and 6, dactylus with unguis not defined; (5)
pleopod 1 digitiform in male with two apical setae,
and subtriangular in female with one seta: (6)
uropod 1 has a ventral peduncular row of stout
setae (not reported in any other ingolfiellidean);
and (7) uropod 2 without basofacial hook on
peduncle, and with five obliquely transverse rows
of setae (not complex setae).

ACKNOWLEDGEMENTS

We thank Ronald Vonk, Zoological Museum of
Amsterdam, The Netherlands for comments on a
first draft of this paper.

REFERENCES

Dahl, E. (1977). The amphipod functional model and its
bearing upon systematics and phylogeny. Z oologica
Scripta 6: 221-228.

Dojiri, M. and Sieg, J. (1987). Ingolfiella fuscina, new
species (Crustacea: Amphipoda) from the Gulf of
Mexico and the Atlantic coast of North America, and
partial redescription of I. atlantisi Mills, 1967.

80

S. Gallego Martinez, G.C.B. Poore

Proceedings of the Biological Society of Washington 100:
494-505.

Lowry, J.K. and Poore, G.C.B. (1989). First ingolfiellids
from the southwest Pacific (Crustacea: Amphipoda)
with a discussion of their systematics. Proceedings of
the Biological Society of Washington 102: 933-946.

Ruffo, S. and Taglianti, A.V. (1989). Description of a new
cavemicolous Ingolfiella species from Sardinia, with
remarks on the systematics of the genus (Crustacea,
Amphipoda, Ingolfiellidae). Annali del Museo Civico di
Storia Naturale Giacomo Doria, Genova 87: 237-261.

Stock, J. (1976). A new member of the crustacean
suborder Ingolfiellidea from Bonaire, with a review
of the entire suborder. Studies on the Fauna of Curagao
and other Caribbean Islands 50: 56-75.

Stock, J. (1977). The zoogeography of the crustacean
suborder Ingolfiellidea with descriptions of new West
Indian taxa. Studies on the Fauna of Curagao and other
Caribbean Islands 55: 131-146.

Stock, J. (1979). New data on taxonomy and
zoogeography of ingolfiellid Crustacea. Bijdragen tot
de Dierkunde 45: 181-190.

Manuscript received January 9 2003; accepted 18 June 2003

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Records of the Western Australian Museum
Volume 22 Part 1 2003

CONTENTS

N. Warburton, C. Wood, C. Lloyd, S. Song and P. Withers
The 3-dimensional anatomy of the North-Western Marsupial Mole
(. Noton/ctes caurinus Thomas 1920) using computed tomography.

X-ray and magnetic resonance imaging 1

F. E. Wells and H. Morrison

Description of Volutoconus hargreavesi calcarelliformis subsp. nov.

(Mollusca: Volutidae) from northwestern Australia 9

R. L. Hoffman

A new genus and species of trigoniuline milliped from Western Australia
(Spirobolida: Pachybolidae: Trigoniulinae) 17

I. Bartsch

Psammophilous halacarids (Halacaridae, Acari) from Dampier,

Western Australia. Description of species and faunal comparison of the
mesopsammal halacarid fauna of western and eastern Australia 23

I. Maryanto

Taxonomic status of the ricefield rat Rattus argentiventer (Robinson and

Kloss, 1916) (Rodentia) from Thailand, Malaysia and Indonesia based on

morphological variation 47

G. C.B. Poore and W.F. Humphreys

Second species of Mangkurtu (Spelaeogriphacea) from north-western Australia 67

S. Gallego Martinez and G.C.B. Poore

A new species of ingolfiellid amphipod (Crustacea: Amphipoda)

from Western Australia 75

Dates of Publication

Records of the Western Australian Museum

Volume 21, Part 1

1 May 2002

Volume 21, Part 2

18 June 2002

Volume 21, Part 3

20 November 2002

Volume 21, Part 4

27 June 2003